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THE INSECTS OF HAWAII

The Insects of Hawaii Series was conceived as a comprehensive taxonomic treatment of theHawaiian insect fauna. With more than 5,000 native arthropod species already described and manymore awaiting discovery, this vast undertaking requires the collaborative efforts of systematists specializing in all components of the class. We have created an editorial board that reflects this diversity ofexpertise. Board members will develop and recruit new projects for the series and will reviewthe scientific content of manuscripts submitted. In view of the efforts required to reach the goal ofcompletely describing Hawaii's entomofauna, the.board invites inquiries from specialists interestedin preparing monographs for groups in need of revision.

Editor: James K. Liebherr Department of EntomologyComstock HallCornell UniversityIthaca, NY 14853-0901

Taxonomic Editors:Rosemary G. Gillespie-University of California, Berkeley (Arachnida)Dan A. Polhemus-Smithsonian Institution, Washington, D.C. (Hemiptera)Manfred Asche-Humboldt Museum fur Naturkunde, Berlin (Homoptera)James K. Liebherr-Cornell University, Ithaca (Coleoptera)Scott E. Miller-Smithsonian Institution, Washington, D.C. (Lepidoptera)Neal L. Evenhuis-Bernice P. Bishop Museum, Honolulu (Diptera) .David B. Wahl-American Entomological Institute, Gainesville (Hymenoptera)

Instructions to Authors

Beginning with the publication of Volume 16, the Insects of Hawaii Series is being produced as acamera-ready publication. Authors are responsible for providing final copy to the publisher,following a style and format consistent with Volumes 16 and 17. Prospective authors should request detailed information on the preparation of camera-ready copy from the editor. For reviewpurposes, contributions should be submitted to the editor in standard double-spaced manuscriptformat.

While there are no page charges for publications in this series, a modest subsidy will be requiredof each author. The amount of this subsidy will be determined by the size and complexity of themanuscript.

Front cover: Hylaeus mana Magnacca and Daly, n. sp.Back cover: Hylaeus difficilis (Perkins) on Dodonaea

I N S E C T S O F H AWA I I

INSECTS OF HAWAII

A Manual of the Insects of the Hawaiian Islands, including an Enumerationof the Species and Notes on Their Origin, Distribution, Hosts, Parasites, etc.

VOLUME 17

HAWAIIAN HYLAEUS (NESOPROSOPIS) BEES

(HYMENOPTERA: APOIDEA)

by HOWELL V. DALY

Professor Emeritus of Entomology

University of California

Berkeley, CA

and KARL N. MAGNACCA

Department of Entomology

Cornell University

Ithaca, NY

University of Hawai‘i Press Honolulu

© 2003 University of Hawai'i Press

All rights reservedPrinted in the United States of America

03 04 05 06 07 08 5 4 3 2 1

library of Congress Cataloging-in-Pub1ication Data(Revised for vol. 17)

Insects of Hawaii; a manual of the insects of the Hawaiian Islands.

Vols. 1-9 by Elwood C. Zimmerman.Vols. 10-14 by D. Elmo Hardy and others.

Vol. 15 by Kenneth Christiansen and Peter Bellinger.

Vol. 16 by James K. Liebherr and Elwood C. Zimmerman.Includes bibliographies and indexes.

Vol. 17 by Howell V. Daly and Karl N. Magnacca

1. Insects-Hawaii. I. Daly, Howell V. II. Magnacca, Karl N.

QL489.H3Z5 595.709969 48-45482ISBN 0-8248-2674-4 (v. 17)

Camera-ready text prepared by the authors.

University of Hawai'i Press books are printed on acid-free

paper and meet the guidelines for permanence and durabilityof the Council on Library Resources.

Printed by The Maple-Vail Book Manufacturing Group

CONTENTS

ABSTRACT ................................................................................................................... 1

INTRODUCTION ........................................................................................................ 3

THE HAWAIIAN ISLANDS ............................................................................... 3

HISTORY OF COLLECTIONS AND TAXONOMY ........................................ 3

DISPERSAL AND EVOLUTION OF HYLAEUS SPECIES IN HAWAII ........ 7

BIOLOGY OF HYLAEUS ................................................................................... 9

NATIVE AND ADVENTIVE ENEMIES .......................................................... 9

ECOLOGY OF HYLAEUS IN HAWAII ........................................................... 11

NATIVE AND ADVENTIVE BEES OF THE HAWAIIAN ISLANDS .......... 12

DESCRIPTIVE TERMINOLOGY .......................................................................... 14

KEYS TO SPECIES ................................................................................................... 24

KEY TO MALES.............................................................................................. 25

KEY TO FEMALES ......................................................................................... 34

TAXONOMY OF HYLAEUS (NESOPROSOPIS) SPECIES ................................ 43

NOMINA DUBIA ..................................................................................................... 209

ACKNOWLEDGMENTS ....................................................................................... 210

BIBLIOGRAPHY .................................................................................................... 211

APPENDIX: RECORDS OF RECENT COLLECTIONS ................................... 217

INDEX ....................................................................................................................... 230

HYLAEUS SPECIES-GROUP NAMES ......................................................... 230

ARTHROPOD ENEMIES AND ASSOCIATES ............................................ 231

PLANTS USED AS NESTING SUBSTRATES ............................................ 231

FLOWER VISITATION RECORDS .............................................................. 231

HAWAIIAN HYLAEUS 1

ABSTRACT

The native colletid bee species of the genus Hylaeus (Nesoprosopis) are revised for theHawaiian Islands based on adult morphology. Sixty species are recognized. NEW SPECIESdescribed by Magnacca and Daly are H. akoko and H. gliddenae from Hawaii; H. kukuiand H. takumiae from Maui; H. kuakea, H. mana, H. mimicus, and H. nalo from Oahu;and H. kokeensis and H. solaris from Kauai. NEW SYNONYMIES are proposed (seniorsynonyms first): H. assimulans (Perkins) = H. assimulans oahuensis (Perkins); H. coniceps(Blackburn) = H. comes (Perkins); H. connectens (Perkins) = H. koae (Perkins), H. insulaeMeade-Waldo (earlier replacement name for Nesoprosopis vicina Perkins), and Prosopistrigona Warncke (unwarranted replacement name for Nesoprosopis vicina Perkins); H.flavipes (F. Smith) = H. blackburni (F. Smith); H. fuscipennis (F. Smith) = H. fuscipennisobscuripes (Perkins), H. fuscipennis swezeyi (Cockerell), and H. caeruleipennis (Perkins);H. haleakalae (Perkins) = H. neglectus (Perkins); H. nivicola (Meade-Waldo) = Prosopisfarinosa Warncke (unwarranted replacement name for Nesoprosopis nivalis Perkins); H.paradoxicus (Perkins) = H. erythrodemas (Perkins); H. rugulosus (Perkins) = Prosopisovatula Warncke (unwarranted replacement name for Nesoprosopis rugulosa Perkins); H.specularis (Perkins) = H. homoeochromus (Perkins); H. unicus (Perkins) = H. binominatusMeade-Waldo (earlier replacement name for Nesoprosopis laticeps Perkins) and Prosopisavara Warncke (unwarranted replacement name for Nesoprosopis laticeps Perkins). NEWCOMBINATIONS are Hylaeus muranus (Warncke) and Hylaeus niloticus (Warncke). Thefollowing names are considered nomina dubia: “Nesoprosopis rubrocaudatus Blackburnand Cameron” and “Nesoprosopis attripennis Perkins.”

The history of collections of the bees, their taxonomy, attributes for dispersal, and theirbiology and ecology are discussed. Taxa of adventive bees are characterized as an aid toidentification of native species. Terminology used in the treatment is explained, followedby keys to the identification of species based on adult males and females. Treatment ofeach species includes annotated synonymy and other references; diagnosis of identifyingfeatures and general distribution; description of male and female; localities where firstcollected and recent collections; flower records; remarks on taxonomic problems and otherinformation; and line illustrations. Recent collection records are provided in an appendix.

INSECTS OF HAWAII VOL. 172

Fig. 1. Map of major islands in the Hawaiian Island chain (after Zimmerman 1948:10).

HAWAIIAN HYLAEUS 3

INTRODUCTION

The native bees of the Hawaiian Islands are believed to have evolved from a single lineageof the genus Hylaeus (family Colletidae) and are assigned to the subgenus Nesoprosopis.The genus Hylaeus is worldwide in distribution with 46 subgenera and an estimated 579species (Michener 2000:62). The subgenus Nesoprosopis is among the largest of thesubgenera with 60 of its species found in the Hawaiian Islands and eight species in theOriental region. The latter includes one species also known from the Palearctic region(Hirashima 1977, Ikudome 1989). About three-quarters of the species described from theHawaiian Islands by Perkins (1899, 1910, 1911) have been collected recently (Table 1).Some are still as rare or as abundant as he observed, yet others, formerly abundant, havenot been collected recently. After more than a century, we have little knowledge of theirmating and nesting habits, food preferences, natural enemies, or roles as pollinators ofendemic Hawaiian plants. The purpose of this volume is to encourage future research byfacilitating identification and by documenting the survival of these interesting bees.

The Hawaiian Islands

Hawaii consists of an east-west chain of volcanic islands, each formed as the Pacific platemoves over a stationary hot spot in the Earth’s crust. There are currently six high islands(Fig. 1), along with two smaller ones and a chain of subsided low islands and coral atollsstretching WNW to Midway. Beyond that, the Emperor Seamounts extend nearly toKamchatka. Eruptive activity is almost entirely restricted to the youngest island, Hawaii,which is about 500,000 years old and still growing. The oldest high island at present,Kauai, is about 5.1 million years old, but it is believed that there has been at least one highisland (i.e., one capable of inducing rainfall and thus supporting a diverse biota) continuouslysince Kure, about 29 million years (Carson and Clague 1995). As islands subside anderode, volcanoes that were previously joined by land bridges are separated by channels.The islands of Maui, Molokai, Lanai, and Kahoolawe (referred to collectively as MauiNui) were once joined in this way; earlier, Molokai was probably joined to Oahu. However,Kauai was never linked to Oahu, nor Hawaii to Maui, although the channels were muchnarrower in the past.

Because of Hawaii’s great isolation – the Islands are over 3500 km from any continentallandmass, the most isolated high islands in the world – the flora and fauna are derived fromrelatively few introductions. As a result, they are depauperate at the family level, but containa number of extraordinary radiations, such as the celebrated Drosophila flies, and theClermontia and Cyanea lobelioids. Hylaeus may be included among these; there are morespecies of the genus in the Hawaiian Islands than the 48 species in America north of Mexico.

History of Collections and Taxonomy

The first specimens of bees from the Hawaiian Islands were collected during the voyage ofthe British ship Blossom commanded by Captain Frederick William Beechey. The specimens


Species Kauai Oahu Molokai Lanai Maui HawaiiH. akokoH. andrenoidesH. angustulusH. anomalusH. anthracinus1

H. assimulansH. chlorostictusH. conicepsH. connectensH. crabronoidesH. difficilisH. dimidiatusH. dumetorumH. facilisH. filicumH. finitimusH. flavifrons4

H. flavipesH. fuscipennisH. gliddenaeH. haleakalaeH. hilarisH. hirsutulusH. hostilisH. hulaH. inquilinaH. kauaiensisH. kokeensisH. konaH. kuakeaH. kukuiH. laetusH. longicepsH. manaH. mauiensis

Table 1. Checklist and distribution of Hylaeus (Nesoprosopis) in the Hawaiian Islands. Key: =original published Perkins’ records (1899, 1910, 1911) and Perkins’ collection, but not recentlyrecollected; = Perkins’ records and recently collected (after 1989); = new records for islands(from recent collections unless noted); = new species. The last row provides the sum of all speciesand the sum of endemic species recorded from each island.

3

2

2

5

8 8

6

8

7

HAWAIIAN HYLAEUS 5

Species Kauai Oahu Molokai Lanai Maui HawaiiH. melanothrixH. mimicusH. muranusH. mutatusH. naloH. niloticusH. nivicolaH. ombriasH. paradoxicusH. peleH. perkinsianus10

H. perspicuusH. psammobiusH. pubescensH. rugulosusH. satellesH. setosifronsH. simplexH. solarisH. specularisH. sphecodoidesH. takumiaeH. unicusH. volatilisH. volcanicusH. species A12

Total/endemic 14/11 15/5 14/1 15/0 24/5 28/18

1 Hylaeus anthracinus reported from Niihau by Beardsley and Tuthill (1959) is probably H.flavifrons; H. anthracinus reported from Kauai by Hopper and Daly (1995) is H. solaris.2 Recently collected from Kahoolawe (CUIC).3 1913 is most recent collection (BPBM).4 Niihau and Lehua; Hylaeus anthracinus reported from Niihau by Beardsley and Tuthill (1959)is probably H. flavifrons.5 Recorded as H. blackburni by Fullaway (1918).6 1934 is most recent record (BPBM).7 Recorded by Fullaway (1918).8 Unpublished records from Perkins’ collection.9 1914 is most recent record (BPBM).10 Nihoa; Timberlake (1926), Beardsley (1966).11 Female only, identification tentative; 1972 is most recent record (BPBM).12 Identifiable as a distinct species, but insufficient material for formal description.

9

11


were labeled “Sandwich I. Beechey” and were presumably taken on Oahu by the ship’snaturalist, George Tradescant Lay. Captain Beechey visited Oahu twice: 19 to 31 May1826, and 25 January to 4 March 1827. On the first visit, Lay arrived ill with dysentery andwas left to recuperate until the ship returned in 1827. Beechey wrote about Lay in hisnarrative (1832:345), “During our absence, he had unfortunately been prevented pursuinghis researches among the islands by a severe illness.” One might assume that Lay wasconfined to Oahu, but evidence exists that Lay or some other member of the expedition didvisit other Hawaiian Islands, including Hawaii (see remarks under species treatment of H.flavipes). Lay did recover and complete the voyage. As a result of the voyage, about halfa dozen species of Hymenoptera from the Hawaiian Islands were deposited in the BritishMuseum (today The Natural History Museum, London). In 1853, the bees were describedas Prosopis anthracina and P. flavipes by Frederick Smith. The genus Prosopis Fabriciuswas later found to be a junior synonym of Hylaeus Fabricius, the generic name now in use(Dathe 1979). Prosopis continues to be a valid subgenus of Hylaeus (Michener 2000:183).

The next collections of bees were made by Reverend Thomas Blackburn, who residedon Oahu from 1876 to 1882 and made brief trips to other islands (Anonymous 1913).From these specimens, also deposited in the British Museum, Smith (1879a, 1879b)described five new species. Shortly afterward, Blackburn, W. F. Kirby, and P. Cameronreviewed the Hawaiian Hymenoptera and described a total of five additional new speciesof bees, bringing the number to twelve (Blackburn and Kirby 1880; Blackburn 1886, 1887).

In 1891, the “Sandwich Islands Committee,” a joint committee of the British Associationfor the Advancement of Science and the Royal Society, selected Robert Cyril Layton Perkinsto investigate the land fauna of the Hawaiian Islands. They could not have made a betterchoice (Scott 1956, Manning 1986, Liebherr and Polhemus 1997, Liebherr and Zimmerman2000). Perkins arrived in Honolulu early in 1892 and worked for the committee until 1902.He continued to live in Hawaii and work as an entomologist until retiring to England in1912. Perkins initially assigned numbers to his collections that were replaced with printedlocality and identification labels (Perkins 1897). The collections from 1892 to 1906 weredivided later and deposited in the Bernice P. Bishop Museum, Honolulu; The Natural HistoryMuseum, London; and the Hope Entomological Collection, Oxford University Museum ofNatural History, Oxford. Locality records for Perkins’ collections from 1892 to 1906 aregiven below under the species treatments.

His monumental Fauna Hawaiiensis includes descriptions of 43 new species-group namesof native bees and a key to the males of 47 species (1899, 1910). Perkins proposed thegenus Nesoprosopis just for the native bees of Hawaii (1899:75), including the 12 speciespreviously described by other authors in Prosopis. In 1911, he added five more new species.Prosopis facilis Smith was designated the type species of Nesoprosopis by Popov(1939:168). Nesoprosopis is currently considered a subgenus of Hylaeus (Michener2000:201).

By an odd coincidence, Perkins on his return to England discovered a bee new to England.The bee was H. pectoralis Forster (=H. kriechbaumeri Forster), which he recognized wasclosely similar to his Hawaiian Nesoprosopis (Perkins 1913:lxxviii). This species is now

HAWAIIAN HYLAEUS 7

known from Europe to Japan (Hirashima 1977:25), but is considered by some authors tobelong to the subgenus Prosopis rather than Nesoprosopis (Dathe 1980).

A large collection of Hawaiian aculeates was made by W. M. Giffard mainly during1901-1915 in which 49 of the 58 described species of native Hylaeus were represented.This provided D. T. Fullaway the opportunity to augment some descriptions and to pointout possible synonyms (1918). The latter apparently have been ignored, probably becauseof his tentative and unconventional statements of synonymy. In our judgment, he was correctin most instances. Giffard’s collections also confirmed the distributions of the speciesrecorded by Perkins. Most locality records for Giffard’s collections (Bishop Museum) andthose of P. H. Timberlake (Los Angeles County Museum) are not provided in this study.

Meade-Waldo (1923) placed Nesoprosopis as a subgenus of Hylaeus and proposed newnames for three homonyms that resulted from his action. The last new taxa to be proposedwere a new species from Nihoa Island by Timberlake (1926) and a new variety by Cockerell(1926). These brought the total to 62 species-group names for the endemic bees of Hawaii.The bees are known from the six main islands plus Kahoolawe, Niihau, and Nihoa. Thesmaller islands are undercollected and may have more species than are recorded fromthem. Daly (1994) designated lectotypes for 44 species-group names and identified holotypesfor 13. Type specimens for five species-group names could not be found.

Recent records reported here date from after 1989 and are based primarily on collectionsfrom all the major islands by K. Magnacca (KM), and from Hawaii and Maui by P. Gambino(PG) and R. Takumi (RT). Due to the large number of localities, only representative sitesare listed under the species description; the full list of complete records and abbreviationsis in the Appendix. Museum collections cited are the British Museum of Natural History(BMNH), Cornell University Insect Collection (CUIC), Bernice Pauahi Bishop Museum(BPBM), Essig Museum of Entomology (UCB), Hawaii State Department of Agriculturecollection (DOA), and Oxford University Museum of Natural History (OUMNH).

Some species, such as H. anthracinus, H. facilis, and H. simplex, were common inPerkins’ time but are now rarely collected. Out of the 60 species recognized here, thefollowing 11 species (18%) have not been recently collected (i.e., since 1989); however,without further searching the species are not presumed to be extinct: H. perkinsianus(Timberlake) from Nihoa; H. finitimus (Perkins) and H. perspicuus (Perkins) from Kauai;H. anomalus (Perkins) and H. nalo new species from Oahu; H. mauiensis (Perkins) and H.melanothrix (Perkins) from Maui; H. gliddenae new species and H. simplex (Perkins)from Hawaii; H. satelles (Blackburn) from Lanai, Maui, and Molokai; and H. niloticus(Warncke) from Molokai, Lanai, and Hawaii.

Dispersal and Evolution of Hylaeus Species in Hawaii

The nearest relatives of the native Hawaiian species are eight Nesoprosopis species inJapan (Hirashima 1977, Ikudome 1989); additional undescribed species may exist in China(Hirashima 1977). However, the geographic origin and characteristics of the initial colonistin Hawaii are unknown. Like some other kinds of bees, Hylaeus have features that shouldfavor dispersal. Female bees are inseminated as young adults and use the stored sperm for


the rest of their lives. The typical longevity of female Hylaeus is unknown. Although malesare usually considered short-lived compared to females, an adult male was kept alive for74 days (Daly and Coville 1982:76). Hylaeus are capable of flight in search of food, mates,and nest sites. Carried by wind to a distant location, an inseminated young female couldmake a nest and produce offspring of both sexes. The nest itself can be transported as well.Some Hylaeus females make their nests in wood or stems that could be carried by waterwith floating vegetation. With these attributes, it is not surprising that native Hylaeus, aswell as adventive species of bees with similar biology, have spread among the HawaiianIslands. However, after reviewing the record, Michener (2000:99) concluded that mostbees are not particularly successful in crossing major barriers. According to Michener, therare colonization of remote oceanic islands is probably limited to small bees such as Hylaeus.

Dispersal between islands appears to be much easier. Unlike many speciose Hawaiiangroups such as Drosophila, about one-third of the known Hylaeus species (21/61, includingH. species A) are not island endemics, including three (H. connectens, H. laetus, and H.specularis) found on all major islands (see Table 1). Only five species each are endemic toOahu and Maui, one to Molokai, and none to Lanai (six of these have been extremelyrarely collected, and they may yet be found on other islands). This indicates that they areprobably able to cross narrow barriers such as those between the islands of Maui Nui withrelative ease. In contrast, most of the species of Kauai (11/14) and Hawaii (18/28) areendemics. Given that Hylaeus are relatively strong fliers and tend to spend time at or abovethe forest canopy rather than below, it is not surprising that they should be carried betweenislands with relative frequency.

Evolutionary relationships among the Hawaiian species are difficult to assess, as manymorphological characters occur in several groups in different combinations. However, fivebasic groups can be relatively easily distinguished. A coastal group, consisting of H.assimulans, H. finitimus, H. flavipes, H. longiceps, H. niloticus, and H. ombrias, ischaracterized by weakly dilated scapes, T6/7 hair golden and appressed or prostrate, facemarks extending above the antennal sockets, and short body hair. Another group, consistingof H. chlorostictus, H. difficilis, H. facilis, H. hirsutulus, H. laetus, H. nivicola, H.perkinsianus, H. simplex, and H. volcanicus, is characterized by an undilated median processof the eighth sternum, T6/7 hair prostrate, and a single central mark on the face, and isprimarily found in dry habitats. Five species, H. hilaris, H. hostilis, H. inquilina, H.sphecodoides, and H. volatilis, are united by parasitic habits and other characters (seebelow). A large group composed of H. angustulus, H. coniceps, H. crabronoides, H.dumetorum, H. filicum, H. kona, H. mana, H. mauiensis, H. melanothrix, H. mimicus, H.pele, H. specularis, and H. species A, is distinguished by three separate face marks, agroove on the underside of the scape, T6/7 hair erect and dark, and a generally smoothpropodeum, and occurs mainly in wet forest. Finally, H. akoko, H. andrenoides, H. anomalus,H. fuscipennis, H. gliddenae, H. kukui, H. muranus, H. paradoxicus, H. perspicuus, H.pubescens, H. satelles, and H. setosifrons form a group of generally large species withpunctured terga. The remaining species possess unusual characters or combinations ofthose that occur in the above groups, and do not fit clearly into one of them.

HAWAIIAN HYLAEUS 9

Biology of Hylaeus

Female bees of the subfamily Hylaeinae, to which Hylaeus belongs, and the AustralianEuryglossinae are unique among bees in carrying all the pollen for their nest provisions intheir internal crop rather than on external scopal hairs (Michener 2000:15). The lack of theusual scopa gives Hylaeus a hairless, wasplike appearance. However, both sexes of Hylaeus(Nesoprosopis) have plumose hairs on the body that are longest on the sides of the thorax.

Females of most species of Hylaeus are believed to construct and provision their ownnests (i.e., they exhibit solitary nesting behavior even when nesting in aggregations). Nestingsites are usually in preexisting cavities such as hollow stems, holes in wood, under bark, increvices or under rocks, and burrows in soil (see plant hosts as nesting substrates in index).Although wood-nesters usually use preexisting beetle and other burrows, sawdust undernest holes of H. pubescens suggests they are capable of enlarging or modifying them.Females can be attracted to trap nests with predrilled holes (Daly and Coville 1982). Thefemale constructs a series of chambers or cells with a cellophane-like membrane producedby the salivary and Dufour’s glands (Espelie et al. 1992). Each cell is provided with pollen,nectar, and an egg. The flowers of native and, occasionally, exotic plants are visited byboth sexes (see flower visitation records in index), but the female alone provisions the nest.Based on observations elsewhere in the world, females of Hylaeus apparently groom thepollen from just their head and forelegs and transfer it to their mouth. The rest of the pollenis discarded (Michener 2000:84). The behavior of both sexes at flowers will be importantto examine in studying the roles of native bees as pollinators of Hawaiian plants. Adventivewasps, ants, and bees such as the honey bee are probably significant competitors for nectar.

Native and Adventive Enemies

Enemies of Hylaeus are poorly known. Five species of native Hylaeus are believed to becleptoparasites of other species of Hylaeus (i.e., the parasitic females enter nests of hostbees and lay their own eggs). Female Hawaiian Hylaeus normally have abundant curvedhairs on the fore tarsi that function in gathering pollen. Perkins (1899:106, 1913:lxxx)discovered five species in which the females have short, straight hairs on the fore tarsi andconsidered them closely allied: H. hilaris, H. hostilis, H. inquilina, H. sphecodoides, andH. volatilis. He associated the straight tarsal hairs with cleptoparasitic habits. If true, thesefive species would be unique among all colletids in their cleptoparasitic behavior. At present,the evidence is circumstantial; no rearing has been done to confirm parasitism. During hisfield studies, KM observed behavior characteristic of cleptoparasites. The bees werecollected almost exclusively while they flew over the ground rather than at flowers. Theybriefly entered nest burrows of other bees and one bee was seen apparently being repelledby the resident. The details of how the cleptoparasitic larva ultimately kills the host egg orlarva and consumes the provisions are unknown. Individuals of the same cleptoparasiticspecies were observed to vary in body size according to the sizes of their presumed host’sprovisions: larger host species provide more food and produce larger cleptoparasites, smaller


host species provide less food (Michener 2000:30). For further discussion of cleptoparasitessee remarks under H. hilaris and H. volatilis in the treatment of species.

The brood also may be attacked by encyrtid and eupelmid parasitoid wasps (see arthropodenemies and associates in index). The most important predators are probably adventiveants, primarily the big-headed ants, Pheidole megacephala (Fabricius) and P. fervens F.Smith; the long-legged ant, Anoplolepis longipes (Jerdon); and the Argentine ant,Linepithema humile (Mayr) (formerly known as Iridomyrmex humilis). These ants areconsidered a serious threat to the endemic arthropod fauna and have extirpated manynative insects from the lowland areas (Reimer 1993, and references therein). AlthoughPerkins (1913:cii) considered adult Hymenoptera to be least susceptible to attack byPheidole, the brood is undoubtedly more vulnerable to these aggressive predators. Pheidolewill even take over honey bee apiaries, killing or driving out the bees and feeding on thebrood (Illingworth 1916).

Both Pheidole and Anoplolepis are primarily restricted to low elevations, while theArgentine ant inhabits montane areas. The absence of Hylaeus from some apparently well-suited coastal habitats may be due to the presence of ants. Argentine ants have beendocumented to severely impact Hylaeus on Haleakala, with no nesting activity found inareas inhabited by ants (Cole et al. 1992). Although the ants are now in relatively smallpockets on Haleakala, if they were to spread over the entire area they could potentiallywipe out the densest population of Hylaeus in the Islands. Because they form very largecolonies and are attracted to sugar, Argentine ants may also be major competitors fornectar.

Adult Hylaeus pubescens may be hosts to the adventive, phoretic mite Afrocypholaelapsafricana (Evans) (Daly and Coville 1982). Some bees are heavily infested, but the natureof injury, if any, is unknown. Other species, including H. fuscipennis, H. paradoxicus, andthe much smaller H. specularis, may also host mites. Gambino (1992) made an analysis ofthe diet of the adventive yellowjacket wasp Vespula pensylvanica (Saussure), a generalistpredator, at natural areas in Hawaii Volcanoes and Haleakala National Parks. He foundadult Hylaeus (Nesoprosopis) sp. among prey items collected by wasp foragers. Althoughthe bees were common at four study sites, they made up only a small fraction of prey items(2 of the 170 arthropods identified to genus) and were retrieved at only one of the sites(Haleakala), suggesting that Hylaeus are not preferred prey.

Perkins (1912) noted the remarkable black body color and dark wings of many Hawaiianaculeates. He considered such wasps and bees conspicuous among day-flying insects. Groupsof wasps and bees were identified that were similar in color patterns and occurred together,yet were from unrelated lineages. The stomachs of many birds were examined and only asingle wasp was found. These color groups of species could readily be explained as examplesof Mullerian mimicry. However, Perkins questioned this interpretation, pointing out thatother bees and wasps, not members of the predominant color group, flew in the sameplaces and were equally ignored by birds. Small Nesoprosopis with clear wings, for example,have been extraordinarily successful. He argued that perhaps it was the general appearanceof an aculeate that was avoided. This left him unable to offer a satisfactory explanation for

HAWAIIAN HYLAEUS 11

the color groups unless they were Mullerian mimics that evolved in response to predatorsnow extinct.

Ecology of Hylaeus in Hawaii

Perkins (1912:688) considered Nesoprosopis to be “almost the most ubiquitous of anyHawaiian insects,” and indeed they range through virtually all native habitats. They can befound from the coastal strand to above 3000 m in the mountains, and from the driest to thewettest locales. A few species are strictly confined to a single habitat type, but most occuracross a range of habitats, though many favor one or another. Nesting habit is a strongerconstraint on habitat suitability than food plant, because ground-nesting species such as H.nivicola require relatively dry areas, while wood-nesters such as H. pubescens can inhabitwetter areas. Either species may sometimes be found foraging in both habitats, because insome areas the wet/dry transition is quite abrupt. Furthermore, some common host plants,notably Metrosideros, occur in both.

Hawaii is noted for its highly diverse plant lineages consisting of rare or localized species,but plant visitation by Hylaeus is largely limited to community-dominant plants. Theseconsist of a small number of species, most of which are members of genera with one or afew representatives in Hawaii: Metrosideros polymorpha (ohia, Myrtaceae), Cheirodendrontrigynum (olapa, Araliaceae), Claoxylon sandwicense (poola, Euphorbiaceae), Acacia koa,Sesbania tomentosa, and Sophora chrysophylla (koa, ohai, and mamane, Fabaceae),Myoporum sandwicense (naio, Myoporaceae), Dodonaea viscosa (aalii, Sapindaceae),Styphelia tameiameiae (pukiawe, Epacridaceae), Scaevola spp. (naupaka, Goodeniaceae),Jacquemontia ovalifolia (pau o Hiiaka, Convolvulaceae), and Sida fallax (ilima, Malvaceae).The one major exception is Chamaesyce (akoko, Euphorbiaceae); Hylaeus visit membersof this genus ranging from the prostrate creeper C. celastroides to the arborescent C.olowaluana. Some other, more diversified groups are among secondary hosts, includingDubautia/Argyroxiphium/Wilkesia and Bidens (Asteraceae). Several of these, notablyDodonaea and Chamaesyce, have very small, nearly petalless flowers that do not conformto the stereotypical “bee flower.” Dodonaea is dioecious, with the male flowers consistingof little more than anthers. While Hylaeus use it as a major pollen source, they have neverbeen observed visiting the female flowers and may not pollinate them. Furthermore, becausethe flowers have little if any nectar, they are rarely visited by male bees. Very few introducedplants are visited, even those with close relatives in Hawaii such as Acacia, with the resultthat Hylaeus are almost completely absent from exotic-dominated habitats. The onlyexception to this rule is the frequent use of Tournefortia argentea (Boraginaceae) by coastalspecies, which may be due to a decline in abundance of the related native Heliotropium.

Flower records are listed under each species. Because only one species of most plantgenera is used, only the genus name of the species listed above is given. Exotic plants areindicated with an asterisk (*). Bees were frequently caught flying around plants (especiallymales searching for foraging females); thus, these records do not necessarily mean that thebee was actually visiting the flower. However, pollen from all plants listed above has beenidentified in crop contents.


Native and Adventive Bees of the Hawaiian Islands

The bees of the Hawaiian Islands include the native bees of the genus Hylaeus (Nesoprosopis)as well as over a dozen other species that arrived after the Islands were colonized byhumans. Here we briefly review the bee fauna as an aid to identifying the native species.The nonnative bees are being treated in detail by R. R. Snelling (in preparation). All beescan be distinguished from wasps by the possession of at least some branched or plumosehair on the body. The hair of wasps is simple or unbranched. The plumose hair of Hylaeusis most conspicuous on the sides of the thorax. Hylaeus are recognizable as small to medium-sized bees (forewing lengths about 3-8 mm), slender bodied, usually black, relatively hairless,with short bilobed tongues, and two submarginal cells in the forewing. Males of mostspecies and females of some species have yellow marks on the face, hence the commonname “yellow-faced bees.” All native species of Hawaiian Hylaeus belong to the subgenusNesoprosopis, characterized by the shapes of the last three metasomal sterna of males: S6,the last visible sternum, has an apical, median emargination; S7, concealed in the genitalchamber, has reduced hemisternites with sparse or no hairs; and S8, also concealed (the tipmay be exposed), has a long, narrow, median process that may be curved and rodlike ordorsoventrally dilated and apically bifid with a fringe of hair.

Two adventive Hylaeus are present in Hawaii. They belong to subgenera other thanNesoprosopis, and both can be distinguished from the native species by the appearance ofthe propodeum in dorsal view: the dorsal anterior area is deeply sculptured (areolate) andseparated sharply from the vertical, flat, posterior area by an irregular, transverse carina.The anterior area of the propodeum of the native Nesoprosopis is usually weakly tomoderately rugose, rarely areolate, and not separated from the rounded, posterior area bya carina. Males also can be separated by the appearance of S6, which is not emarginate inthe adventive species. Of the two adventive species, H. (Hylaeus) bisinuatus Forster probablycame to North America from Europe and then to Hawaii (Beardsley 1959:23, Snelling1975:8). This species is small (forewing length about 3.5 mm) with a black body, yellowmarkings, and coarse, dense punctation. It has been collected only twice on Oahu, in 1959and 1994, and may not be established. Recently, a second species was discovered, theAustralian H. (Gnathoprosopis) albonitens (Cockerell). This distinctive small bee (forewinglength about 3 mm) has a dark metallic blue body and extensive ivory markings on the faceand legs (Houston 1981:52, Hopper and Daly 1995:35). It has been collected from allislands except Maui and Lanai. It is usually found on ohia in dry lowlands where nativebees are absent, but occasionally overlaps with them at middle elevations.

The only other bees in the Hawaiian Islands with two submarginal cells in the forewingare the Megachilidae, which are distinguished by their long tongues and thick, hairy bodies.More likely to be mistaken for Hylaeus are two species of small carpenter bees, Ceratinaarizonensis and C. (Neoceratina) sp. nr. C. dentipes Friese. The former has been collectedon Oahu and the Kona coast of Hawaii; the latter is widespread on all the major islands andappears to be parthenogenic, as only females have been collected. These are small bees(forewing length about 2.5-3 mm) that are black, relatively hairless, with facial and other

HAWAIIAN HYLAEUS 13

marks, but have long tongues and three submarginal cells in the forewing. The facial markof the female, a single small median ivory spot, is unlike that of any native Hylaeus.

Hylaeus is a member of the large family Colletidae. Colletids are among the families ofbees that are informally called “short-tongued” because the tongue is relatively short inproportion to body size. In these bees the blunt mouthparts and short-segmented palpi arevisible projecting forward beneath the head or retracted and partly concealed behind themandibles. Adventive short-tongued bees in Hawaii are the two Hylaeus species mentionedabove, and a species of sweat bee (Halictidae), Lasioglossum (Dialictus) impavidum(Sandhouse), which are small (forewing length about 3 mm), dull metallic green bees withthree submarginal cells in the forewing.

The other major group of bees, the “long-tongued” bees, have slender, elongatemouthparts thrust forward or folded back under the head. These include species in Hawaiisuch as the honey bee, Apis mellifera L. (Apidae) with pollen baskets on the hind legs; sixspecies of Megachile and one Lithurgus (Megachilidae), with two submarginal cells in theforewing; small carpenter bees, genus Ceratina (Apidae), including the two black speciesalready mentioned, and a bright metallic green species, Ceratina (Pithitis) smaragdula(Fabricius); and a large carpenter bee, Xylocopa (Neoxylocopa) sonorina F. Smith (Apidae),a very large bee with black females and yellow-brown males. All species appear to beadventive with the exception of Apis, which was purposely introduced in 1857 (Crane1990:27).

Most adventive bees are found primarily in nonnative habitats and are not majorcompetitors with the native Hylaeus. However, Ceratina are often found in large numbersin native coastal habitats and may have an impact on Hylaeus populations. The impact maynot be entirely detrimental. Ceratina excavate nest burrows in dead twigs, a behavior thatmight create nest sites for Hylaeus. Ceratina smaragdula, discovered in 1995 on Oahu,has built up a substantial population at Kaena Point, the only recent collection site forcoastal Hylaeus on the island. It now makes up a large proportion of visits to Sesbaniatomentosa, an endangered shrub, and is expanding its range (D. Hopper, pers. comm.). Inother areas, Apis is a major competitor for nectar and pollen resources and is often veryabundant in native habitats.


DESCRIPTIVE TERMINOLOGY

Terms used in this present text and frequently encountered in previous literature on Hylaeus(Nesoprosopis) are given here. The main references on terminology used here are Eady(1968), Harris (1979), and Nichols (1989). Illustrations of the anatomy of bees can befound in Michener (2000). More detail on the anatomy of Hylaeus in particular may befound in Houston (1975). The descriptions of species are intended to include the variationobserved in the available specimens. These are often few in number and may be old anddiscolored. Many species that were rarely collected in Perkins’ time remain rare in collections.Specimens collected during the last four years have been used whenever possible. Variationbeyond that described here will no doubt be found as more specimens are examined. Thisis to be expected, especially for the extent of markings, wing color, punctation, and propodealsculpture.

As indications of dimensions of various features of the male head, distances were measuredto the nearest 0.02 mm at ×64 (Fig. 2C-E) and expressed here as millimeters ×100. Themeasurements are based on one specimen, usually a type or specimen from Perkins’ collectingefforts. Some quantification, however imperfect, is also required to express size andproportions. Ratios are based on too few specimens to provide useful statistics, so therange of values is given and must be considered approximate. The dilation of the scape,length of the malar space, and width of the supraclypeal area are expressed as ratios (furtherexplanation below). As an indication of body size, the length of the forewing, from thebase of the costal vein (Fig. 3) to the apex, was measured to the nearest 0.1 mm. Bodysizes of bees are divided into three categories: small (forewing length less than about 4mm), medium-sized (about 4-6 mm), and large (more than about 6 mm).

Colors. We have used simple terms for body colors, such as yellow, red, brown, orblack. Where an area or structure is of an intermediate color it may be described as red-brown or yellow-brown. Colors of the wing membranes are clear, smoky, brown, or darkbrown. The membrane may also be iridescent. The cuticle of specimens collected by Perkinsoften has faded to a pale brown or brownish black in comparison with fresh specimens thatare black.

Cuticle. The general sclerotized cuticle of the exoskeleton, used here in reference to thegeneral melanized color (yellow-brown, brown, or black) of a leg or other body part incontrast to yellow or ivory marks.

Face. The face is divided by several fine lines called sulci (Snodgrass 1956:18; sutures ofMichener 2000:40). The face also has topographic areas that are approximately delimitedby imaginary lines rather than by sulci (Fig. 2A-B). The vertex is the area on the top of thehead surrounding the ocelli. The frons is the area immediately below the vertex and abovethe clypeus, including the area between and above the antennal sockets. The frons iscontinuous with the upper paraocular areas at each side and delimited by imaginary linesrunning between the lateral ocelli and antennal sockets. That part of the frons below andbetween the antennal sockets is usually called the supraclypeal area. It is defined at eachside by the subantennal sulci and apically by the frontoclypeal sulcus (median part of theepistomal sulcus). The shape of the supraclypeal area is expressed as a ratio of the width

HAWAIIAN HYLAEUS 15

Fig. 2. Face of a male Hylaeus bee. A-B. Features of the face: a, acme (median prominence betweenantennal sockets); cl, clypeus; es, epistomal sulcus; fc, frontoclypeal sulcus (median part of epistomalsulcus); fl, frontal line; fr, frons; ia, impunctate areas above antennal sockets; lp, lower paraoculararea; mf, middle of frons; ms, malar space; sc, supraclypeal area; ss, subantennal sulcus; tp, tentorialpits; up, upper paraocular area; v, vertex. C-E. Measurements: Malar space, t; eye, a: b: c: d; clypeus,e × f; frons, g × h: i; interalveolar, j: k : l: m, D (diameter) n; interocellar, o: p: q: r, D (diameter) s.

Cr

q

a

j k

d

b

gh

E

e

f

ilm

n

op

s

tD

cc

up

lp sc

cl

fr

esms

up

lp

A

mf

a

ia fl

tp

ss

v

B

fc


(frontoclypeal sulcus)/length (subantennal sulcus from outer rim of antennal socket tojunction with frontoclypeal suture). A longitudinal groove or frontal line is sometimesvisible at the midline below the median ocellus. A narrow linear area just at each side of themidline or frontal line is here called the middle area and is often distinctively punctured.The prominence between the antennal sockets is defined here as the frons acme. Theparaocular area extends along the sides of the face parallel to the eye. The lowerparaocular area is at or below the antennal sockets and is separated from the clypeus bythe epistomal and subantennal sulci. The upper paraocular area is above the antennalsockets and separated from the frons by an imaginary line between the lateral ocellus andantennal socket. In females, a narrow groove, the facial fovea, runs along the upper marginof the eye and may terminate dorsad below or above an imaginary, transverse line tangentto the anterior edges of the lateral ocelli. The paraocular area, mainly the lower area, maybe visibly depressed relative to the surface of the eye and clypeus.

Frontal, lateral, and caudal views. The frontal view of a body part is seen by theobserver in front of the specimen. The frontal view of the head is the view from immediatelyin front of the bee with the plane of the bee’s face at right angles to the long axis of thebody. The caudal view is the view from immediately behind the bee. The frontal view of theforeleg is the view of the leg as if it were extended laterally at a right angle to the long axisof the body, revealing the anterior surface of the leg. Specifications for viewing the scapeare complicated by the various positions that the antennae can assume on specimens. Thescape of males is usually somewhat flattened and may be “dilated” or expanded. The frontalview of the antennal scape is defined here as the view of the scape as if the scape is positionedvertically upward and appressed to the surface of the face, revealing the maximum widthand length of the scape. In this position, the punctured surface of the scape is visiblewhereas the impunctate underside next to the face is hidden underneath. Also in this position,the scape has a “median margin or edge” that is often convex (morphologically dorsal orposterior if the axis of the antenna is parallel to the long axis of the body and rotated so theconvex margin is up) and a “lateral margin or edge” that is usually nearly straight(morphologically ventral or anterior). The median edge of a moderately dilated scape isbroadly rounded (convex) and often with longer hairs, whereas the lateral edge is more orless straight. In species with a strongly dilated scape both edges are convex. The degree ofa scape’s arch is seen from the side in an edge-on view of the lateral margin.

Ground. The ground is the surface of the cuticle excluding punctures (pits) and rugae.Different surfaces that reflect light from the microscope illuminator can be described as:shiny or polished, reflecting the maximum amount of light; lustrous, a bright, but diffusereflection of light; slightly lustrous, a less intense reflection; and dull, the minimum reflectionof light. The usual surface texture can be described by the term coriaceous, which refers toa microscopic fine network of lines, possibly representing the outlines of epidermal cells.Types of coriaceous ground used here are: smooth, showing only the network of fineinterconnecting lines, producing a shiny or lustrous surface; nearly smooth, with each cellraised in a low, round bump; granular, with each “cell” elevated enough to produce arough, less shiny surface; minutely wrinkled, with elevations aligned in short wrinkles to

HAWAIIAN HYLAEUS 17

produce a wavy pattern and slightly lustrous or dull surface; minute parallel wrinkles,elevations aligned in longer, more or less parallel wrinkles, also known as a striate surface.

Malar space. The malar space is the area between the lower eye margin and the base ofthe mandible; also known as the “cheek.” Length is measured as the minimum distancebetween the eye margin and edge of the space next to the base of the mandible. The lengthof the malar space is compared with the diameter of the median ocellus. The diameter ofthe median ocellus (DMO; Fig. 2D, s) is measured as the transverse distance across theclear area of the ocellus. The clear area of the ocellus and not the maximum diameter of thelens itself was chosen because the exact outside, dark edge of the lens is sometimes difficultto see. The ratio “malar length/diameter of median ocellus (DMO)” is used to specify thelength of the malar area: very short (0.2 or less of DMO); short (0.3 to 0.4); medium (0.5to 0.6 DMO or about half the diameter of the median ocellus); long (0.7-1.0 or almost aslong as the diameter of the median ocellus); and extremely long (greater than 1.0 or longerthan the diameter of the median ocellus). The ocelli of females are often smaller than thoseof the males, resulting in a higher ratio for equal malar lengths.

Mark. Marks are limited areas of reddish, yellow, or ivory color surrounded by thegeneral color of the cuticle. Marks may occur on the face, pronotal collar and lobes, tegula,median plate of the forewing (Fig. 3), and legs. Entire segments of the legs may be markedwith a single color. An obscure mark is one off-color (brownish or reddish yellow) andwhose boundaries are diffuse, not sharp. Perkins (1899) used the term vitta (pl., vittae) fora stripe or band of color. See also “pale area” below.

Mesosoma, metasoma, T, S. A bee’s body is divided in three parts like most insects, butwhat appears to be the “thorax” is the thorax combined with the first abdominal segmentor propodeum (see below). This middle subdivision of the body is called the mesosoma. Indorsal view, from front to rear, are the pronotum, scutum, scutellum, metanotum, andpropodeum. In lateral view, the mesepisternum is the large sclerite below the wings andbetween the fore and middle legs. What appears to be the “abdomen” is the second andsucceeding abdominal segments and is called the metasoma. The terga and sterna of the

Fig 3. Base of right fore wing showing the median plate, which is yellow in H. kokeensis and H.kona. The median plate is brown in all other species. Note that length of the forewing is measuredfrom the base of vein C to the apex of the wing.

Edge ofscutum

Median plate

Base of vein CCostal sclerite

Tegula

CR

M+CuV


metasoma are abbreviated and numbered as T1 and S1 for the first metasomal segment andso forth. The disk of the tergum is the central area.

Pale area. As in “pale area” of the fore tibia; an area of cuticle of the fore tibia, andsometimes fore femur, seen in frontal view that is weakly melanized, somewhat transparent,and pale in color, but different from spots or stripes of yellow or ivory pigments that arecalled marks. The pale area is not considered here to be a mark. However, the area may becombined with a yellow spot or stripe along the dorsal margin. The pale area is usuallypresent on males and sometimes on females. Important: For brevity in the diagnoses, thepale area is not mentioned, and a bee may be said to be entirely black in color or unmarkedeven though the bee may have the tibial pale area.

Pronotal lobes and collar. The lobes are the rounded, posterolateral part of the pronotumjust in front of the wing bases; the collar is the narrow, transverse dorsum of the pronotumjust visible behind the head and in front of the scutum. These structures may be markedwith yellow or ivory.

Propodeum. The propodeum is subdivided topographically into the ventrolateral,punctured, and pubescent “sides” and the impunctate, hairless, dorsal basal area of thepropodeum (Michener 2000:45). The sides and basal area may be sharply delimited oronly obscurely so by an impressed line. In Hylaeus (Nesoprosopis) the basal area has abroad, subhorizontal “anterior area” that is variously sculptured. Sculpture “at the front”or “at the base” means sculpture along the very anterior margin of the basal area. Theanterior area is continuous with the posterior, subvertical surface of the propodeum alonga transverse brow that serves as the boundary between the anterior and posterior surfaces.The brow has no discrete landmarks and is merely the rounded transition from subhorizontalto subvertical surfaces. Sculpture “to the brow” means sculpture extended over most ofthe anterior area, from the front to the rounded transition or brow. The basal area continuesbeyond the brow, forming a triangle on the posterior surface. A lateral carina arisesposterolaterally on the propodeum and diminishes anterolaterally; the carina may be strongand discrete or merely a ridge.

Pubescence. Hair is described as simple (without branches), weakly plumose (somehairs with at least one branch and usually mixed with simple hairs that may be in the majority),and plumose (hairs mostly with more than one branch). Plumosity can be difficult to discernwhen branches or the primary hairs are short. The length of hairs on the scutum can beimportant in distinguishing similar species. Because the scutal pubescence typically occursas a dense underlayer with scattered longer hairs, the measurements for it are presented inthe form 8-12 (in millimeters ×100, as for the dimensions of the head). The first numberrepresents the average height of the lower layer, and the second the maximum height of thelonger hairs. Hair length should be measured in lateral view. On the scutum, average hairlength often increases from front to back, and the extreme anterior and posterior areasshould be ignored. For females, the hairs of the vertex are sometimes useful; these includethose on the top of the head just behind the ocellar triangle (hairs within the ocellar trianglemay be differentiated and distinctly longer). In a few wide-ranging species, notably H.flavipes and H. difficilis, specimens from higher elevations have longer hair than thosenear the coast. Hairs are easily rubbed off, and on specimens where this is the case those

HAWAIIAN HYLAEUS 19

left may be shorter than the length given. In all cases, a few unusually long hairs may bepresent. The coloration of the hairs, especially those of the apex of the abdomen, can beuseful. The color must be used with caution because variation in lighting can make thehairs appear lighter or darker. Description of T6 or T7 hairs refers to those on the dorsalarea of the sclerite as seen in lateral view; those along the lateral and apical margins aretypically much longer. The hairs may be described as erect (at least a 45° angle to thecuticle), prostrate (more bent down), or appressed (hairs dense, decumbent, and in closecontact, though not necessarily pressed down on the cuticle). Hairs that are bent over nearthe base are not considered erect even if they arise perpendicular from the cuticle. Thenormal posture of the hairs can be altered by retraction of the abdomen, matting of hairs,or other disturbance, resulting in erect hairs that appear prostrate and vice versa.

Punctation, density of pits. The density of pits is expressed in terms of “pit widths” asestimated by eye. The pits of very dense punctation are less than 1 pit width apart and maybe so close together that the pits are contiguous and distorted or subpolygonal in shape.Less dense punctation is expressed as “1 pit width apart” or “1-2 pit widths apart” and soforth. The density and distinctness of pits should be measured in frontal view at relativelyhigh magnification; even impunctate areas often have micropits that are visible at lowangles.

Rugae. Rugae (sing., ruga) are wrinkles on the surface of the cuticle. As used here forthe sculpture of the basal area of the propodeum, types of rugosity are lineately rugose,rugae more or less straight, parallel, and longitudinal with reference to the body axis;irregularly lineately rugose, rugae that are not interconnecting, but somewhat forked ornot parallel or not longitudinal (oblique or transverse with reference to the body axis);reticulately rugose, interconnecting or netlike; and clathrately rugose, latticed, elevatedridges intersecting at right angles. The last two types may form shiny depressions betweenthe rugae and be described as areolate.

Scape. The basal segment of the antenna is the scape. In frontal view (see definitionabove), the scape of the male may be nearly cylindrical or variously dilated (expanded)and, in lateral view, more or less arched (broadly bent). Degrees of dilation of the scapeare measured by the ratio of width/length. Width is measured as the maximum at a rightangle to the length; length is measured from the apex of the scape to the base just above thebasal constriction. Categories of dilation of the scape are not or weakly dilated (W/L ratio0.4 or less), slightly dilated (0.5-0.6), moderately dilated (0.7-0.8), strongly dilated (0.9-1.0), and extremely dilated (1.0 or more) (Fig. 4A-J). The arch of the scape, seen in anedge-on view of the lateral margin, is estimated by eye and described as not arched, slightlyarched, moderately arched, or strongly arched (Fig. 4K-P). The maximum arch of thescape is the highest point above an imaginary line connecting the base and apex of thescape. The maximum arch may be in the middle or closer to the apex of the scape. Sincethe arch is sometimes diagonal across the scape, it may be necessary to view it at a slightapical angle in order to clearly see the full arch of the lateral edge. The scape of eachspecies is illustrated here in frontal view and lateral view. The lateral-view drawings of thescape for the new species include a line indicating the lateral edge (not always obviouswhen the thickness varies). The impunctate underside (posterior in frontal view) of thescape of some species may have a groove or pit near the median edge. This appears to be


Fig. 4. Scapes of male Hylaeus antennae, showing variation. A-J. Frontal views of bee’s left scapeand pedicel showing degrees of dilation: l, lateral margin on right of figure; m, median margin; widthindicated by dashed lines and W/L ratios in parentheses. F. Shows concave lateral margin and angularlateral apex, called “hooked.” K-P. Lateral views of scape and pedicel showing degrees of arch. L-N.Shows maximum arch near middle. O-P. Shows maximum arch near apex.

C (0.5) E (0.7)D (0.6) F (0.7)A (0.3) B (0.4) Weakly dilated Slightly dilated Moderately dilated

G (0.8) I (1.2) J (1.4)H (1.0) Strongly dilated Extremely dilated

K L O P Not arched Slightly arched

Moderately dilated

M NModerately arched Strongly arched

m l

HAWAIIAN HYLAEUS 21

the outlet for a gland in the scape and may be similar to those reported for Australianhylaeines (Houston, 1975:11).

Tarsus. The basitarsus is the long, basal tarsal segment and the distitarsus is the lasttarsal segment. The tarsal segments in between are the mediotarsus. We call the mediotarsusand distitarsus together the terminal tarsal segments.

Terminalia of male. The term terminalia is applied here to S7 and S8 plus the genitalia,all of which normally are retracted into the metasoma and concealed by T7 and S6. S7 isreduced and divided medially into sclerotized and lightly pigmented, external hemisterniteseach with a lateral portion that may be elongated posteriorly in a lateral process of S7(Fig. 5A). The length of the lateral process may be less than, subequal to, or greater thanthe transverse width of the hemisternite. The lateral process may be described as tapered (avariety of shapes, gradually narrowed from base to apex), parallel-sided (elongate processwith more or less parallel sides and with a rounded apex) or sword-shaped (process relativelybroad throughout its length and with a pointed apex). Internally, each hemisternite also hasan anterior, elongated lateral apodeme of S7. The hemisternites of S7 and their apodemesfit closely against S8 just anterior to the median process of S8. The hemisternites and theirapodemes are illustrated here in ventral view. Although in life they normally are in a curvedplane, they are shown here as if they were flat to show the maximum outline. S8 has abroad, internal apodemal base with an external, posterior median process (Fig. 5B). Theprocess may be essentially linear with a nearly uniform diameter like a curved rod (Fig.6A-I) or it may be dilated (expanded dorsoventrally) to various degrees. Dilated processesusually have a flexure (obtuse angle in the dorsoventral plane) at midlength. At this pointthe dilation is usually broader, creating a prominent hump. The dilation is a flat, membranouscuticle, usually hairy along the ventroposterior edge. The dilation may be variable withinspecies and on some specimens may be partly collapsed or folded, appearing smaller than

Fig. 5. Terminalia of male Hylaeus. A. Seventh sternum, ventral view: la, lateral apodeme; lp, lateralprocess. B. Eighth sternum, ventral view, and apex, caudal view. C. Eighth sternum, lateral view: ba,basal apodeme; f, flexure of process; mp, median process; t, tip of apical bifurcation. D. Genitalia,dorsal view: gb, gonobase; gf, gonoforceps; pv, penis valves.

A B C D

lp

la

mp f

t

t

gb

gf

pv

t

ba


Fig. 6. Median processes of eighth sternum of male Hylaeus, lateral view, and apex, caudal view,showing variation. A. H. hirsutulus. B. H. mutatus. C. H. difficilis. D. H. laetus. E. H. nivicola. F. H.volcanicus. G. H. simplex. H. H. facilis. I. H. chlorostictus. J. H. anthracinus (two forms). K. H.kuakea. L. H. psammobius. M. H. ombrias. N. H. andrenoides. O. H. sphecodoides. P. H. flavipes.Q. H. connectens. R. H. kauaiensis. S. H. kokeensis. T. H. dumetorum. U. H. muranus. Extent ofpubescence shown by dashed lines.

E F G H I

J K L M N O

A B C D

P Q R S T U

HAWAIIAN HYLAEUS 23

normal. Degrees of dilation are: weakly dilated, a slight dorsoventral expansion (Fig. 6J-N); moderately dilated, a distinct, but narrow membrane beneath the process (Fig. 6O-P);and strongly dilated, a broad membrane beneath the process (Fig. 6Q-U). The apex of theprocess may be simple (no bifurcation) or bifurcate with tips of various sizes and shapes:filamentous, straplike, or broad, round tips. The process and tips are usually pubescent, insome species so much so that the extent of dilation and the shape of the cuticular tipsare obscured by matted hair. Care should be taken in distinguishing cuticle from hair becausethe descriptions refer to the shape of the cuticular structures. The entire S8 is shown herein a ventral view with the tip shown in a caudal view (dorsal side up) and also in a lateralview (Fig. 5C). The male genitalia has a basal sclerite, the gonobase, an inner pair ofnarrow penis valves (sagittae of Perkins 1910), and an outer pair of broad gonoforceps(stipites) that are apically pubescent (Fig. 5D). The apices of the gonoforceps may besubequal to the apices of the penis valves or, if the gonoforceps are expanded apically,exceed them; only in a few species are the gonoforceps shorter. The genitalia are shownhere in dorsal view.


KEYS TO SPECIES

The keys are designed for identification and do not necessarily indicate cladistic relationships.Characteristics of adventive species of bees and an explanation of anatomical terms andmeasurements are given in the preceding section. In each couplet, the characters given firstare easiest to see or most reliable. Read through the entire couplet before deciding, assome have qualifications or exceptions. Some species appear more than once in the keys toinclude intraspecific variation. Maui Nui refers collectively to the islands of Maui, Lanai,and Molokai.

To identify males, it may be necessary to measure certain structures with an ocularmicrometer and to partly extract the male terminalia. Couplet 2 separates species with themedian process of S8 dilated and bifurcated with broader tips (Fig. 6M-U), versus theprocess not dilated and with very slender or no tips (Fig. 6A-I). For most specimens thedecision is obvious, but some specimens may appear to be intermediate, especially if thetips are shriveled or the dilation of the process is collapsed. If in doubt, the best choice is toassume the process is dilated (i.e., couplet 2a). Five of the potentially troublesome species(Fig. 6J-L) have been keyed under both choices. The process of H. anthracinus is variablein width; H. kuakea, H. mana, H. nalo, and H. psammobius have weak dilations.

Females are more difficult to key because they have fewer structural and color characters.Many common species are far down in the key, and often characters are not as obvious orwell defined as in the males. Their identification is greatly aided by narrowing downpossibilities using associated males. It is not uncommon, however, to collect mainly malesof one species and mainly females of another at a given site, especially when the totalnumber of specimens is low. Males and females may also frequent different species ofplants in the same area.

Nearly all females can be separated, though some couplets (notably 4, 5, and 50) mayrequire experience with multiple species to identify them correctly. Females of H. difficilis,H. facilis, and H. simplex cannot be reliably distinguished. However, the first is the mostcommon and widespread species on Hawaii and Maui Nui; the latter two, although formerlyabundant, have become extremely rare since Perkins’ time. Thus, unless an associatedmale is available, specimens can generally be assumed to be H. difficilis. Females should becollected carefully to keep them clean and dry. Avoid damaging the pubescence, positionthe antennae forward so that the upper frons is exposed, and pry open the mandibles sothat the apical teeth are visible.

Correct association of males and females is often difficult in Hylaeus due to the generalizedmorphology of the latter. Most species for which recently collected males and females areavailable have been associated using DNA sequences (KM, unpubl. data). These total 39species; a further 10 can be unambiguously associated due to unique characters shared byboth sexes. Of the remaining species, 4 can be placed with relative certainty based on closeassociation with and similarity to males, or by process of elimination. Finally, H. finitimus(only type female known) and H. perkinsianus (two females known, but no specimens areavailable) are provisionally placed in the key as our best guess based on earlier descriptions

HAWAIIAN HYLAEUS 25

and our notes. Hylaeus akoko, H. gliddenae, H. kuakea, and H. nalo were not includedbecause the females are unknown. The female of H. akoko would probably key to H.pubescens, and that of H. gliddenae to H. paradoxicus. The female of H. mauiensis couldnot be included because the original description is insufficient and no specimen is available.

Males have the antenna with 13 segments; metasoma with 7 visible terga; scape of antenna(long basal segment) often, but not always, dilated; hairy structures may project from apexof metasoma; no sting; face and legs often, but not always, marked with yellow.

Females have the antenna with 12 segments; metasoma with 6 visible terga; scape ofantenna subcylindrical, never dilated; slender, hairless sting may project from apex ofmetasoma; face and legs often, but not always, unmarked.

Key to males

1. Scape weakly to moderately dilated in frontal view (Fig. 4A-G; W/L ratio up toabout 0.8); or if about as wide as long, then paraocular marks are present ..................................................................................................................... 2

— Scape strongly to extremely dilated, always as wide as long or wider than long(Fig. 4H-J); paraocular area black .............................................................. 5

2(1). Median process of S8 weakly to strongly dilated (Fig. 6J-U) and with the apexbifurcate with hairy tips of various sizes and shapes, tips sometimes long andstraplike but not very slender or filamentous .............................................. 3

— Median process of S8 not dilated, narrow and linear, of nearly uniform diameterthroughout, and with the apex simple or bifurcate with very slender orfilamentous tips much longer than wide (Fig. 6A-I) ................................... 8

3(2). Basal area of propodeum lineate or reticulately rugose over entire anterior areato brow; T7 hair variable ............................................................................ 4

— Basal area of propodeum mostly smooth coriaceous, rugae at most reachinghalfway to brow, often with only a few very short rugae at the anterior margin;T7 hair erect, dark brown to black ........................................................... 22

4(3). Metasomal terga distinctly punctured (at least T3-7; may be somewhat shallow insmall specimens, but still clear); vertex and scutum hair long, (16-)20-30 ormore and 10-20 respectively ..................................................................... 39

— Metasoma impunctate, pits (if any) very shallow and indistinct; vertex hair shorter,4-20(-24); scutum hair often short, 4-20 .................................................. 48

5(1). Median process of S8 very broadly dilated, almost quadrate in lateral view (Fig.6U) .............................................................................................................. 6

— Median process of S8 only slightly dilated, with a distinct flexure or arc ......... 7


6(5). Scape wider than long, oval in profile (Fig. 46E; W/L ratio about 1.5); clypeusdark or with an irregular ivory mark; sculpture of basal area of propodeumvarying from weakly lineate at base to reticulately rugose over most of thebasal area; Hawaii ................................................................................................................... Hylaeus muranus (Warncke), new combination (p. 145)

— Scape width nearly equal to length, circular in profile (Fig. 59E; W/L ratio about1.1); clypeus dark or, sometimes together with lower frons, of a yellowishred color; basal area of propodeum sparsely sculptured with linear rugae atthe base, if at all; Maui Nui .............. Hylaeus satelles (Blackburn) (p. 179)

7(5). Metasoma conspicuously metallic blue; clypeus with a more or less parallel-sidedivory mark (Fig. 11E); Kauai .......... Hylaeus andrenoides (Perkins) (p. 45)

— Metasoma entirely dark brown to black; clypeus unmarked or with a small subapicalyellow mark; Maui ....................... Hylaeus takumiae, new species (p. 195)

8(2). Malar space long (ratio 0.8-1.2 DMO) .............................................................. 9— Malar space short (ratio 0.2-0.4 DMO) ........................................................... 10

9(8). Malar space very long (ratio about 1.2 DMO); paraocular depression deep andwith a rounded ridge below the antennal socket; scape moderately dilated(W/L ratio about 0.7); Maui ........ Hylaeus nivicola Meade-Waldo (p. 155)

— Malar space long (ratio about 0.8 DMO), paraocular depression shallow andwithout a ridge below the antennal socket; scape only weakly to slightlydilated (W/L ratio about 0.4-0.5); Hawaii, Maui .................................................................................................. Hylaeus volcanicus (Perkins) (p. 205)

10(8). Face with paraocular marks adjacent to eye (sometimes filling entire area betweeneye and clypeus; Figs. 47E, 56E); clypeal yellow mark, if present, partly orentirely separated from paraocular yellow marks by dark areas along sulciand usually with no yellow mark on supraclypeal area ............................. 11

— Face with a single yellow mark, often including supraclypeal area; paraocularmarks, if present, adjacent to clypeal sulci and relatively narrow (Figs.14E,50E) ........................................................................................................... 14

11(10). Paraocular depression deep, evident in frontal view; punctation of scutum dense,pits about 1 pit width apart over most of the disc .................................... 12

— Paraocular depression shallow or absent, appearing flat in frontal view; punctationof scutum not particularly dense, pits 2-3 pit widths apart ...................... 13

12(11). Paraocular mark small, near eye, clypeus black; gonoforceps blunt, shorter thanpenis valves (Fig. 48); Oahu ... Hylaeus nalo, new species, in part (p. 150)

— Paraocular mark nearly or entirely filling in area below antennal socket, clypeusalmost entirely yellow; gonoforceps slightly pointed, longer than penis valves(Fig. 56); Hawaii, Maui .... Hylaeus psammobius (Perkins), in part (p. 170)

HAWAIIAN HYLAEUS 27

13(11). Face almost entirely yellow, paraocular marks extending to or above antennalsockets; all tibiae with yellow markings; median process of S8 with apexbifurcate (Fig. 42B, E); Oahu ..................................................................................................................... Hylaeus mana, new species, in part (p. 135)

— Face with apical clypeal mark and narrow paraocular marks not extending aboveantennal sockets; tibiae black except for pale area; median process of S8simple, somewhat spatulate (Fig. 47B, E); Kauai ...................................................................................................... Hylaeus mutatus (Perkins) (p. 148)

14(10). Scape moderately to strongly dilated (Figs. 14E, 38E; W/L ratio 0.7-0.9), clearlywider in middle than at apex; median process of S8 weakly dilated, compressedlaterally in cross-section (Fig. 6J-K) ......................................................... 15

— Scape weakly to slightly dilated (Fig. 20E; W/L ratio 0.4-0.6), widest point nearapex; median process of S8 rodlike, round in cross section at midlength (Fig.6A-I) .......................................................................................................... 16

15(14). Hair of vertex and scutum short (6-10 and 4-6 respectively); frons with pits notmuch closer medially than laterally; paraocular mark usually wide (Fig. 14E);Hawaii, Maui Nui, Oahu .... Hylaeus anthracinus (F. Smith), in part (p. 52)

— Hair of vertex long (16-24), of scutum medium (8-12); frons with pits densemedially, nearly contiguous; paraocular mark usually narrow (Fig. 38E); Oahu............................................ Hylaeus kuakea, new species, in part (p. 125)

16(14). Gonoforceps not expanded apically or medially, subequal to the penis valves inlength and not contiguous (Fig. 7C-D) ..................................................... 17

— Gonoforceps expanded apically so that they project beyond the penis valves andmay touch medially (Fig. 7A-B) ............................................................... 18

17(16). Median process of S8 with apex simple or bifurcate with short tips, apex alwaysspatulate (Fig. 20B); scutum hair longer (12-20), usually reddish brown;Hawaii, Maui Nui .................................. Hylaeus difficilis (Perkins) (p. 71)

— Median process of S8 with apex bifurcate with long, slender tips (Fig. 40B);scutum hair shorter (8-12), white to pale brown; Hawaii, Maui Nui, Oahu,Kauai ....................................................... Hylaeus laetus (Perkins) (p. 129)

18(16). Lateral process of S7 very short, triangular (Fig. 7F); median process of S8 simple;scape slightly arched (Fig. 31B, F) ........................................................... 19

— Lateral process of S7 long and broad (Fig. 7E); median process of S8 bifurcatewith long slender tips; scape moderately arched (Fig. 23B, F) ................ 20

19(18). Hind tibia with large basal yellow mark; wings smoky; Kauai ....................................................................................... Hylaeus hirsutulus (Perkins) (p. 106)

— Hind tibia unmarked or with only obscure yellow spots at base; wings clear; Nihoa............................................... Hylaeus perkinsianus (Timberlake) (p. 166)


20(18). Legs mostly unmarked or with small obscure spots of yellow, no basal white oryellow stripe on hind tibia ......................................................................... 21

— Legs marked with yellow, including basal yellow stripe on hind tibia; Kauai ............................................................. Hylaeus chlorostictus (Perkins) (p. 58)

21(20). Gonoforceps with pubescence very short or missing at inner apex (Fig. 23D);Maui Nui, Oahu ...................................... Hylaeus facilis (F. Smith) (p. 80)

— Gonoforceps with pubescence extended around apex to inner angle (Fig. 61D);Hawaii ................................................... Hylaeus simplex (Perkins) (p. 184)

22(3). Underside of scape deeply arched, much more so than upper side (Figs. 35F, 45F)and/or apex abruptly thickened in lateral view, almost hooked (Figs. 44F,66F) ........................................................................................................... 23

— Scape gently arched, may be gradually thickened toward the apex but not with anabrupt projection (e.g., Fig. 36F) ............................................................. 27

23(22). Paraocular marks extending well above antennal sockets ............................... 24— Paraocular marks not extending above antennal sockets ................................. 25

24(23). Scutum hair medium (6-8); apex of scape gradually thickened in lateral view;pronotum and bases of mid and hind tibiae clearly marked with yellow; processof S8 with tips two or more times longer than wide (Fig. 45B); Oahu .............................................................. Hylaeus mimicus, new species (p. 142)

— Scutum hair long (12-20); apex of scape abruptly thickened in lateral view;pronotum and tibiae unmarked or bases of tibiae with only yellowish traces;process of S8 with tips twice as long as wide or less than twice (Fig. 24B);Hawaii ......................................................Hylaeus filicum (Perkins) (p. 83)

Fig. 7. Genitalia of male Hylaeus, dorsal view (A-D) and seventh sternum, ventral view (E-F) showingvariation. A and E. H. facilis. B and F. H. hirsutulus. C. H. difficilis. D. H. laetus.

A B C D

E F

HAWAIIAN HYLAEUS 29

25(23). Supraclypeal area distinctly wider than long, W/L ratio 1.5-2.5 ..................... 26— Supraclypeal area about as wide as long; Maui ...................................................

...................................................... Hylaeus melanothrix (Perkins) (p. 140)

26(25). Clypeus entirely yellow or nearly so, paraocular marks short and narrow, not orbarely extended above clypeal mark; process of S8 without a flexure atmidlength (Fig. 35C, E); Kauai ........ Hylaeus kauaiensis (Perkins) (p. 117)

— Clypeus black or with a subapical yellow spot up to about half its length; paraocularmarks, if present, longer than clypeal mark or extended to antennae; processof S8 with a flexure at midlength (Fig. 66C, E); Maui Nui, Oahu .................................................................. Hylaeus unicus (Perkins), in part (p. 198)

27(22). Median plate at base of forewing distinctly yellow (Fig. 3); or if only an obscureorange color, mandible with a large yellow mark ..................................... 28

— Median plate brown; mandible without markings (except sometimes H.crabronoides with a small, obscure mark) ................................................ 29

28(27). Legs extensively marked with yellow, all tibiae with large basal and smaller apicalmarks often joined by a dorsal stripe; mandible with a yellow mark; scapewith a distinct groove on the underside of the median edge, greatest widthnear middle, often marked with yellow; Hawaii ........................................................................................................ Hylaeus kona (Blackburn) (p. 122)

— Legs not extensively marked, hind tibia at most with a yellow mark extended halfits length; mandible black; scape without a distinct groove underneath, greatestwidth near apex, unmarked; Kauai ................................................................................................................... Hylaeus kokeensis, new species (p. 119)

29(27). Scape not marked with yellow ......................................................................... 30— Scape marked with yellow along lateral edge; Hawaii .........................................

................................................................... Hylaeus hula (Perkins) (p. 112)

30(29). Median process of S8 moderately dilated, at least basally, usually with a distincthump (Figs. 19C, 21C) ............................................................................. 31

— Median process of S8 hardly dilated, of even width in lateral view, without ahump (Fig. 42C); Oahu ......... Hylaeus mana, new species, in part (p. 135)

31(30). Scape nearly flat (Fig. 12F) .............................................................................. 32— Scape slightly but distinctly arched (Fig. 19F) ................................................. 33

32(31). Upper frons with a distinct patch of black hairs, conspicuous in frontal and lateralview; Hawaii .....................................Hylaeus dumetorum (Perkins) (p. 78)

— Upper frons with a dark area of dense sculpture but not a conspicuous patch ofblack hairs; Maui and Lanai .............. Hylaeus angustulus (Perkins) (p. 47)


33(31). Paraocular marks longer against eyes than clypeus (Fig. 17E) or filling in entirearea up to antennal sockets (Fig. 19E); or if absent, the clypeal mark issubapical and extends less than half the length of the plate, or absent ..... 34

— Paraocular marks entirely adjacent to clypeus (as in Fig. 20E) or absent; clypeusall yellow or with a large central mark (Fig. 18E); Hawaii, Maui Nui, Oahu,Kauai .....................................Hylaeus connectens (Perkins), in part (p. 64)

34(33). Clypeus usually partly to mostly yellow, clypeus basal margins and supraclypealarea always black (Figs. 17E, 19E); frons with dark patch of hair or densersculpture in middle; scape with a small pit or groove on the underside of themedian edge (sometimes indistinct) .......................................................... 35

— Clypeus entirely yellow (Fig. 21E); frons without dark patch of hair in middle,sculpture even; scape without a groove on the underside of the median edge;Hawaii ................................................ Hylaeus dimidiatus (Perkins) (p. 75)

35(34). Clypeus with a yellow mark, usually large; if reduced, then paraocular marks arereduced or absent ...................................................................................... 36

— Clypeus black (known only from type; individuals with a small apical mark maybe found), paraocular marks present; Maui ............................................................................................................ Hylaeus mauiensis (Perkins) (p. 138)

36(35). Scutum hairs mostly short, 4-8, with only scattered longer hairs to 14; paraocularmarks large, filling in area up to antennal sockets or nearly so ................ 37

— Scutum hairs long, 14-18; paraocular marks usually small or absent, not reachingantennal sockets; Hawaii, Maui ................................................................................................................. Hylaeus coniceps (Blackburn), in part (p. 61)

37(36). Head in dorsal view normal, less than half as long as wide; frons with impunctate,shining areas above antennal sockets and laterad of medial dark patch ... 38

— Head in dorsal view thick, about half as long (from acme to middle of occiput) aswide (may be difficult to distinguish in isolation, but obvious in comparisonwith any other species); frons punctate over entire area; Hawaii ........................................................................ Hylaeus crabronoides (Perkins) (p. 68)

38(37). Flagellum of antenna yellow underneath; pronotal collar unmarked, lobessometimes with small marks; Hawaii ......... Hylaeus pele (Perkins) (p. 163)

— Flagellum of antenna dark underneath; pronotal collar and usually lobes markedwith yellow; Hawaii, Maui?, Oahu, Kauai ............................................................................................................. Hylaeus specularis (Perkins) (p. 189)

39(4). Pronotal collar with or without a narrow transverse ivory stripe, median breadthof stripe, if present, less than diameter of median ocellus; tibiae less extensivelymarked with ivory ..................................................................................... 40

— Pronotal collar with very broad transverse ivory stripe; full length of all tibiaemarked with ivory; Kauai ................ Hylaeus perspicuus (Perkins) (p. 168)

HAWAIIAN HYLAEUS 31

40(39). Metasoma entirely red ...................................................................................... 41— Metasoma dark brown to black, or red only on T1 ......................................... 42

41(40). Paraocular area, clypeus, and scape marked with yellow; Hawaii ............................................................................. Hylaeus gliddenae, new species (p. 98)

— Face entirely black; Hawaii .................. Hylaeus paradoxicus (Perkins) (p. 160)

42(40). Scape with a distinct stripe along the full length of the lateral edge ................ 43— Scape unmarked, entirely orange-brown (H. anomalus), or with an obscure orange

mark near the base of the lateral edge (H. kukui) ..................................... 45

43(42). Scape with a narrow stripe along the lateral edge, which is distinctly convex; palearea only on front tibia .............................................................................. 44

— Scape broadly ivory on the lateral third, lateral edge not distinctly convex; paleareas on front and mid tibiae, and all femora; Hawaii ............................................................................................... Hylaeus akoko new species (p. 43)

44(43). Second metasomal sternum with subapical hump visible in lateral view; Hawaii......................................................... Hylaeus pubescens (Perkins) (p. 173)

— Second metasomal sternum evenly convex, without a distinct hump; Maui Nui,Oahu ................................................ Hylaeus fuscipennis (F. Smith) (p. 94)

45(42). Lateral margin of scape straight or nearly so; paraocular area usually marked;paraocular depression shallow .................................................................. 46

— Lateral margin of scape distinctly convex; paraocular area unmarked; paraoculardepression deep, shining; Maui .......... Hylaeus kukui, new species (p. 127)

46(45). Scape black; legs black, usually with yellow or ivory marks ........................... 47— Scape and legs largely orange-brown; Oahu ........................................................

............................................................ Hylaeus anomalus (Perkins) (p. 50)

47(46). Face with three yellow marks (Fig. 29E), or entirely yellow; Maui, Molokai ............................................................. Hylaeus haleakalae (Perkins) (p. 100)

— Paraocular areas nearly entirely filled in with ivory, clypeus unmarked or withonly a narrow, transverse, apical band (Fig. 60E); Hawaii ....................................................................................Hylaeus setosifrons (Perkins) (p. 181)

48(4). Paraocular depressions deep, distinct in frontal view ...................................... 49— Paraocular depressions shallow or absent ........................................................ 51

49(48). Paraocular mark small or absent, clypeus black; gonoforceps blunt, shorter thanpenis valves (Figs. 48D, 58D) ................................................................... 50

— Paraocular mark nearly or entirely filling in area below antennal socket, clypeusalmost entirely yellow; gonoforceps slightly pointed, longer than penis valves(Fig. 56); Hawaii, Maui .... Hylaeus psammobius (Perkins), in part (p. 170)


50(49). Median process of S8 very weakly dilated, without a hump at flexure; gonoforcepsrounded apically (Fig. 48); Oahu .................................................................................................................. Hylaeus nalo, new species, in part (p. 150)

— Median process of S8 strongly dilated, with a distinct hump; gonoforcepssomewhat truncate apically, with a distinct angle medially (Fig. 58); Hawaii......................................................... Hylaeus rugulosus (Perkins), (p. 176)

51(48). Paraocular marks not extended above antennal sockets, or absent; clypeus entirelyyellow or with only narrow margins black (Figs. 18E, 25E) .................... 52

— Paraocular marks extended in a stripe up to or above antennal sockets (Figs.15E, 26E); or if reduced or absent, then clypeus with a small spot or unmarked(Fig. 17E) .................................................................................................. 55

52(51). Median process of S8 moderately dilated, with a pronounced hump (Figs. 18C,25C) .......................................................................................................... 53

— Median process of S8 weakly dilated, hump absent or poorly developed (Figs.14C, 38C) .................................................................................................. 54

53(52). Scape moderately dilated, W/L ratio 0.6-0.7; paraocular marks narrow or absent(Fig. 18E); Hawaii, Maui Nui, Oahu, Kauai ...............................................................................................Hylaeus connectens (Perkins), in part (p. 64)

— Scape strongly dilated (W/L ratio 0.8-1.0); paraocular marks broad, usually fillingarea below antennal socket (Fig. 25E); Kauai, Niihau, Lehua .................................................................................... Hylaeus flavifrons (Kirby) (p. 87)

54(52). Hair of vertex and scutum short (6-10 and 2-6 respectively); frons with pits notmuch closer medially than laterally; paraocular mark usually relatively broad(Fig. 14E); Hawaii, Maui Nui, Oahu ....................................................................................................... Hylaeus anthracinus (F. Smith), in part (p. 52)

— Hair of vertex long (16-24), of scutum medium (8-12); frons with pits densemedially, nearly contiguous; paraocular mark usually narrow (Fig. 38E); Oahu............................................ Hylaeus kuakea, new species, in part (p. 125)

55(51). Scape with lateral edge nearly straight in frontal view, or very slightly and evenlyconcave ..................................................................................................... 56

— Scape with lateral edge distinctly concave in frontal view, projecting somewhatat apex (Fig. 34E); Hawaii ................. Hylaeus inquilina (Perkins) (p. 114)

56(55). Face long and narrow, supraclypeal area distinctly longer than wide (W/L ratio0.6-0.8; Figs. 41E, 62E) ........................................................................... 57

— Face normal, supraclypeal area about as long as wide ..................................... 58

HAWAIIAN HYLAEUS 33

57(56). Scape weakly dilated (W/L ratio 0.3-0.4), median edge more or less straight,greatest width near apex (Fig. 41E); Maui Nui, Oahu ........................................................................................... Hylaeus longiceps (Perkins) (p. 133)

— Scape moderately dilated (W/L ratio 0.7-0.8), median edge strongly convex,greatest width near middle (Fig. 62E); Kauai ........................................................................................................ Hylaeus solaris, new species (p. 187)

58(56). T7 hair short, prostrate or appressed, plumose, golden to brown ................... 59— T7 hair long, erect, weakly plumose, dark brown to black; Hawaii, Maui ..........

............................................. Hylaeus coniceps (Blackburn), in part (p. 61)

59(58). Median process of S8 with bifurcations short and relatively broad (Fig. 67B);lateral carina of front coxa lamellate, drawn at base into a distinct angle or ashort spine (Fig. 8D; may be weakly developed in H. hostilis) ................ 60

— Median process of S8 with bifurcations narrow and straplike (Fig. 15B); lateralcarina of front coxa not lamellate or spinose (Fig. 8C) ............................ 62

60(59). Yellow mark on face extended to or above antennal sockets in a narrowing andpointed stripe (Fig. 67E) or reduced; scape unmarked; red color, if present,only on basal segments of metasoma ........................................................ 61

— Yellow mark on face extended well above antennal sockets in a broad stripe (Fig.32E); scape marked with yellow or orange; basal segments of metasomaalways orange or red, often with extensive reddish patches on mesosoma;Kauai ..................................................... Hylaeus hostilis (Perkins) (p. 109)

61(60). Malar space short (ratio about 0.3 DMO); Maui Nui, Oahu .......................................................................................... Hylaeus volatilis (F. Smith) (p. 201)

— Malar space very short (ratio about 0.2 DMO); Hawaii .......................................................................................... Hylaeus sphecodoides (Perkins) (p. 192)

62(59). Metasoma entirely black, without hair bands; scape black or with a yellow mark,but never entirely yellow ........................................................................... 63

— Basal segments of metasoma red, apical segments usually with white hair bands(sometimes rubbed off); scape entirely yellow; Maui Nui .......................................................................................... Hylaeus hilaris (F. Smith) (p. 103)

63(62). Median process of S8 moderately dilated, with a pronounced hump (Fig. 26C);all tibiae usually with extensive yellow markings; face with yellow paraocularmark usually extended as a broad stripe above antennal sockets (Fig. 26E;specimens from higher elevations sometimes with a narrowing stripe) ... 64

— Median process of S8 slightly dilated, hump weak or absent (Figs. 15C, 49C);mid and hind tibiae entirely black except for small basal marks; face withparaocular mark extended as a narrowing stripe above the antennal sockets(Figs. 15E, 49E) ........................................................................................ 65


64(63). Hind tibiae mostly yellow, at least basal half and apical ring marked; Hawaii,Maui, Lanai ........................................... Hylaeus flavipes (F. Smith) (p. 90)

— Hind tibiae mostly black, with yellow marks only at the very base and apex; Kauai(known only from description and key by Perkins 1899:100, 1910:602) .................................................................. Hylaeus finitimus (Perkins) (p. 86)

65(63). Scape with greatest width near middle, about twice as long as wide (W/L ratioabout 0.5), hair on median edge plumose; clypeal mark often reduced ... 66

— Scape with greatest width near apex, more than twice as long as wide (W/L ratioabout 0.4), hair on median edge mostly simple; clypeus always entirely yellow;Hawaii, Lanai, Molokai ........................................................................................................... Hylaeus niloticus (Warncke), new combination (p. 152)

66(65). Wings slightly smoky to smoky; face with more yellow marking: clypeus entirelyyellow or nearly so, supraclypeal area with or without a yellow mark; Maui,Lanai, Oahu ....................................... Hylaeus assimulans (Perkins) (p. 55)

— Wings dark smoky; face with less yellow marking: clypeus with a large centralspot of yellow or no mark, supraclypeal area without a yellow mark; Hawaii............................................................. Hylaeus ombrias (Perkins) (p. 158)

Key to females

1. Setae of front mediotarsus erect, especially ventrally, distinctly and rather abruptlycurved at apex (Fig. 8A); no projection on lateral edge of front coxa, carinavery narrow and indistinct or absent (Fig. 8C) ........................................... 2

— Setae of front mediotarsus straight or gently arcing, prostrate, similar to othertarsal setae (Fig. 8B); front coxa sometimes with a lateral lamellate carinaterminating proximally in a projection or spine (Fig. 8D) .......................... 6

2(1). Younger islands .................................................................................................. 3— Kauai or Northwestern Islands ......................................................................... 10

3(2). Metasomal terga impunctate or with very minute, indistinct pits; usually smallerspecies, wing length about 6 mm or less; basal area of propodeum usuallymore or less lineate to weakly reticulate or mostly smooth ....................... 4

— Metasomal terga distinctly punctured, at least on T3-6; basal area of propodeumreticulate (except H. satelles); usually large species, wing length about 6 mmor greater (except H. haleakalae) ............................................................ 23

4(3). Basal area of propodeum roughened all over, the rugae sometimes weak but notsmoothly coriaceous anywhere on the dorsal surface; or if smooth near brow,then T6 hairs prostrate, pale brown ............................................................ 5

— Basal area of propodeum usually with very short lineate rugae at front, some-times extending about halfway to brow, but brow beyond rugae alwayssmoothly coriaceous; T6 hairs erect, usually dark brown to black .......... 31

HAWAIIAN HYLAEUS 35

Fig. 8. Left fore tarsi (posterodorsal view) and fore coxae (frontal view) of female Hylaeus. A. H.difficilis, showing curved, pollen collecting setae. B. H. volatilis, a cleptoparasite, showing straightsetae. C. H. inquilina, showing narrow carina along lateral edge. D. H. sphecodoides, frontal viewof fore coxa showing broad lamellate carina (base of carina marked by dashed lines).

0.5 mm

A

B

C D


5(4). Punctures of upper frons clearly evident but shallow, without distinct edges,blending into ground in frontal view; T6 hairs pale to brown, short and/orprostrate .................................................................................................... 46

— Punctures of upper frons distinct in frontal view, relatively deep (may be obscuredby sculpture of ground); T6 hairs variable: golden and prostrate, or dark anderect ........................................................................................................... 50

6(1). Hawaii ................................................................................................................. 7— Older islands ....................................................................................................... 8

7(6). Metasoma and legs entirely black; coxal lamella absent or very narrow, without aprojection (Fig. 8C); Hawaii .............. Hylaeus inquilina (Perkins) (p. 114)

— At least T1 and/or mid and hind tarsi reddish brown; front coxa with a distinctlateral lamella and a dorsal projection (Fig. 8D); Hawaii ................................................................................. Hylaeus sphecodoides (Perkins) (p. 192)

8(6). Red marks sometimes present on metasoma and face but not mesosoma; MauiNui and Oahu .............................................................................................. 9

— Extensive red markings over entire body; Kauai ............................................................................................................... Hylaeus hostilis (Perkins) (p. 109)

9(8). Apical terga of metasoma with white hair bands (may be indistinct or lost byabrasion); coast of Maui Nui ................ Hylaeus hilaris (F. Smith) (p. 103)

— Metasoma lacking hair bands; coast and mountains of Maui Nui and Oahu .................................................................. Hylaeus volatilis (F. Smith) (p. 201)

10(2). Paraocular marks present ................................................................................. 11— Paraocular area unmarked ................................................................................ 14

11(10). Median plate of forewing dark brown to black; clypeal mark absent, or if present,transverse and apical ................................................................................. 12

— Median plate of forewing yellow (Fig. 3); clypeus with at least a small longitudinalmark; Kauai ................................. Hylaeus kokeensis, new species (p. 119)

12(11). Basal area of propodeum with very short rugae at front; with or without markson prothoracic lobes; scutum hairs short, 2-8 .......................................... 13

— Basal area of propodeum with distinct, widely spaced, lineate rugae, extendingabout halfway to brow; without marks on prothoracic lobes; scutum hairsmedium, basal layer about 6 with longer hairs 12-20; Kauai ................................................................................. Hylaeus kauaiensis (Perkins) (p. 117)

HAWAIIAN HYLAEUS 37

13(12). Punctation of upper frons distinct, medially denser with pits less than 1 pit widthapart; scutum very dull; Hawaii, Maui?, Oahu, Kauai ............................................................................... Hylaeus specularis (Perkins), in part (p. 189)

— Punctation of upper frons shallow, pits about 1 pit width apart even around medianline; scutum slightly lustrous; Kauai .... Hylaeus mutatus (Perkins) (p. 148)

14(10). Cuticle without metallic blue tint or metasomal hair bands ............................. 15— Body, especially metasoma, somewhat metallic blue; apical terga of metasoma

with hair bands; Kauai ..................... Hylaeus andrenoides (Perkins) (p. 45)

15(14). Prothoracic collar unmarked ............................................................................ 16— Prothoracic collar with a broad, solid stripe; Kauai .............................................

........................................................ Hylaeus perspicuus (Perkins) (p. 168)

16(15). Mandible normal, with two teeth (Fig. 9A) ..................................................... 17— Mandible with three subequal apical teeth (Fig. 9B); Kauai, Niihau, Lehua .......

............................................................... Hylaeus flavifrons (Kirby) (p. 87)

Fig. 9. Right mandibles of female Hylaeus. A. H. longiceps, showing typical form with two apicalteeth (grooves marked by dashed lines). B. H. anthracinus, with three apical teeth. C. H. rugulosus,with no apical teeth and broad apex.

0.5 mm

A

B C


17(16). Hairs of upper frons and vertex short, 4-8 ....................................................... 18— Hairs of upper frons and usually vertex medium to long, 10-14 or more ........ 19

18(17). Upper frons with pits evenly spaced; Kauai .................................................................................................................. Hylaeus solaris, new species (p. 187)

— Upper frons with pits denser medially; Kauai .................................................................................................................... Hylaeus finitimus (Perkins) (p. 86)

19(17). Scutum hairs short, mostly 4-6 with a few up to 12; vertex hairs medium, 6-16.................................................................................................................. 20

— Scutum hairs longer, 8-16; vertex hairs long, 14-30 ........................................ 21

20(19). T6 hairs long, dark brown to black, erect; Hawaii, Maui Nui, Oahu, Kauai .....................................................Hylaeus connectens (Perkins), in part (p. 64)

— T6 hairs short, pale, prostrate; Hawaii, Maui Nui, Oahu, Kauai ............................................................................. Hylaeus laetus (Perkins), in part (p. 129)

21(19). Clypeus without a longitudinal line .................................................................. 22— Clypeus with a longitudinal depression or faint impunctate line (may be faint or

difficult to see; most prominent just below the frontoclypeal suture, bestobserved from an oblique frontodorsal view); Kauai .......................................................................................... Hylaeus chlorostictus (Perkins) (p. 58)

22(21). Wings dark smoky; Kauai ......................... Hylaeus hirsutulus (Perkins) (p. 106)— Wings subhyaline; Nihoa ............... Hylaeus perkinsianus (Timberlake) (p. 166)

23(3). Cuticle of at least part of metasoma and/or head (other than clypeus) red ..... 24— Cuticle almost entirely dark brown to black .................................................... 25

24(23). Entire metasoma red, head black; Hawaii .................................................................................................................. Hylaeus paradoxicus (Perkins) (p. 160)

— Head and first segment of metasoma red; Oahu .............................................................................................................. Hylaeus anomalus (Perkins) (p. 50)

25(23). Clypeal hairs pale to red-brown, prostrate or decumbent, inconspicuous; hindtibia with a small mark (up to about half the length) or none .................. 26

— Clypeus with conspicuous erect, black hairs; hind tibia with an ivory stripe theentire length; Hawaii ........................Hylaeus setosifrons (Perkins) (p. 181)

26(25). Hawaii ............................................................................................................... 27— Maui Nui or Oahu ............................................................................................ 28

HAWAIIAN HYLAEUS 39

27(26). Metasoma with faint blue reflections; S2 normal; pronotum unmarked; T6 hairsblack, erect; Hawaii ................................................................................................................. Hylaeus muranus (Warncke), new combination (p. 145)

— Metasoma without blue reflections; S2 with a more or less distinct medianprotuberance; pronotal collar with or without an ivory stripe; T6 hairs reddish,densely plumose; Hawaii .................. Hylaeus pubescens (Perkins) (p. 173)

28(26). Punctation of frons not particularly dense, pits not crowded and distorted even atmiddle; scutum and vertex hairs pale, white to yellow; punctation of T2-3strong; larger species, wing length usually over 7 mm ............................. 29

— Punctation of frons very dense, pits crowded and distorted even laterally; scutumand vertex hairs dark brown to black (may appear pale due to reflection oflight); punctation of T2-3 not very strong; smaller species, wing length usuallyless than 6.5 mm........................................................................................ 30

29(28). Pronotum unmarked; face with or without reddish or yellow marks; Maui Nui .......................................................... Hylaeus satelles (Blackburn) (p. 179)

— Pronotal collar with an ivory stripe; face black; Maui Nui, Oahu................................................................................ Hylaeus fuscipennis (F. Smith) (p. 94)

30(28). Hairs of lower paraocular area short, 4-10; face black; Maui ........................................................................................ Hylaeus kukui, new species (p. 127)

— Hairs of lower paraocular area very long, 14-20; sometimes with paraocular marks;Maui and Molokai ............... Hylaeus haleakalae (Perkins), in part (p. 100)

31(4). Paraocular marks large and broad (Fig. 45G), usually extending well above theantennal sockets; or if only slightly above, a longitudinal or triangular clypealmark is usually present .............................................................................. 32

— Paraocular marks small and narrow, not or barely extended above the antennalsockets, or absent; clypeal mark, if present, transverse and apical (Fig. 42G).................................................................................................................. 33

32(31). Median plate yellow (Fig. 3); clypeus entirely black; Hawaii ........................................................................................... Hylaeus kona (Blackburn) (p. 122)

— Median plate brown to black; clypeus with at least a small apical mark; Oahu ...................................................... Hylaeus mimicus, new species (p. 142)

33(31). Supraclypeal area short, W/L ratio 2.5 or more; or if ratio about 2.0, thepropodeum is almost entirely smooth coriaceous and flat with a pronouncedbrow .......................................................................................................... 34

— Supraclypeal area relatively long, W/L ratio about 2.0; propodeum with lineaterugae about halfway to brow; basal area sloping, brow rounded; Hawaii ...................................................................... Hylaeus hula (Perkins) (p. 112)


34(33). Hairs of lower paraocular area long, 10-20 ..................................................... 35— Hairs of lower paraocular area short, 4-10 ...................................................... 36

35(34). Face with paraocular marks; Hawaii ...............Hylaeus filicum (Perkins) (p. 83)Face entirely black; Maui ..................... Hylaeus melanothrix (Perkins) (p. 140)

36(34). Prothorax unmarked, or with marks only on the lobes or collar (not both); facevariable ............................................................................................................................................................................................................................ 37

— Prothoracic collar and lobes both marked; paraocular area usually marked (somespecimens may have facial marks reduced) .............................................. 41

37(36). Basal area of propodeum with dorsal surface not as long as scutellum, and sloping,brow not particularly pronounced; mesosoma convex in lateral view; winglength greater than 4.0 mm ....................................................................... 38

— Basal area of propodeum with horizontal surface long, about as long as scutellum,with brow pronounced; mesosoma depressed, nearly flat in lateral view; smallspecies, wing length usually less than 4.0 mm; Hawaii ................................................................................................... Hylaeus pele (Perkins) (p. 163)

38(37). Vertex and scutum hair medium to long, 10-20 and (4-)6-16 respectively ..... 39— Vertex and scutum hair short, 6-10 and 2-6 respectively; Molokai .....................

.......................................................................... Hylaeus species A (p. 208)

39(38). Pale area of front tibia obscure (sometimes with a small yellow spot at base oftibia); face in lateral view gently convex ................................................... 40

— Pale area of front tibia bright orange or yellow; face in lateral view strongly convex;Maui Nui, Oahu ..................................... Hylaeus unicus (Perkins) (p. 198)

40(39). Punctation of upper frons distinct, relatively deep, pits crowded and distortedmedially; Hawaii, Maui ........ Hylaeus coniceps (Blackburn), in part (p. 61)

— Punctation of upper frons evident but shallow, pits not crowded medially exceptsometimes near frontal line; Hawaii, Maui Nui, Oahu, Kauai ....................................................................Hylaeus connectens (Perkins), in part (p. 64)

41(36). Face in lateral view only slightly protruding beyond the inner margin of the eyes.................................................................................................................. 42

— Face strongly bulging outward, very obvious in lateral view; Hawaii ........................................................................Hylaeus crabronoides (Perkins) (p. 68)

42(41). Upper frons with pits about 1 pit width apart, sometimes slightly closer mediallybut not crowded or distorted anywhere .................................................... 43

— Upper frons with dense punctation, less than 1 pit width apart, pits crowded anddistorted at least near the frontal line ....................................................... 44

HAWAIIAN HYLAEUS 41

43(42). Upper frons smooth, shiny; only known specimen with large paraocular marksand a transverse clypeal mark (Fig. 42G); Oahu ..................................................................................................... Hylaeus mana, new species (p. 135)

— Upper frons minutely granular, slightly lustrous; paraocular marks small, sometimesreduced to an obscure spot, clypeus black; Hawaii ................................................................................................. Hylaeus dimidiatus (Perkins) (p. 75)

44(42). Scutum hairs short, mostly 4-6, longer hairs to 10 common ........................... 45— Scutum hairs extremely short, mostly 2, very few longer hairs; Maui, Lanai ......

.......................................................... Hylaeus angustulus (Perkins) (p. 47)

45(44). Impunctate area laterad of lateral ocellus shiny, surface sculpture faint or indistinct;dull area between median ocellus and antennal sockets broad, about twice aswide as ocellus, and strongly contrasting with lustrous lateral frons; winglength usually 3.8 mm or less; uncommon, local; Hawaii, Maui?, Oahu, Kauai............................................. Hylaeus specularis (Perkins), in part (p. 189)

— Impunctate area laterad of lateral ocellus only slightly lustrous, distinctly coriaceousor minutely granular; dull area between median ocellus and antennal socketsnarrow, only about as wide as median ocellus, and not strongly contrastingwith lateral frons; wing length usually 3.8 mm or more; very common andwidespread; Hawaii ...........................Hylaeus dumetorum (Perkins) (p. 78)

46(5). Scutum hair long, 12-20, usually red-brown ................................................... 47— Scutum hair short, 6-10, white to pale brown; Hawaii, Maui Nui, Oahu, Kauai

.................................................... Hylaeus laetus (Perkins), in part (p. 129)

47(46). Malar area medium to long, ratio greater than 0.6 DMO ................................ 48— Malar area short, ratio 0.6 DMO or less .......................................................... 49

48(47). Malar area very long, ratio greater than 1.0 DMO; longest hairs of lower paraoculararea about 20; Maui ..................... Hylaeus nivicola Meade-Waldo (p. 155)

— Malar area medium length, ratio 0.8-1.0 DMO; longest hairs of lower paraoculararea about 14; Hawaii, Maui ............ Hylaeus volcanicus (Perkins) (p. 205)

49(47). Hawaii, Maui Nui; very common ................. Hylaeus difficilis (Perkins) (p. 71)— Hawaii; rare ..................................................Hylaeus simplex (Perkins) (p. 184)— Maui Nui, Oahu; rare ..................................... Hylaeus facilis (F. Smith) (p. 80)

50(5). T6 hairs conspicuously erect, dark, usually simple or sparsely plumose ......... 51— T6 hairs prostrate or appressed, golden brown, densely plumose ................... 55

51(50). Malar area short, ratio about 0.5 DMO or less ................................................ 52— Malar area long, ratio about 0.8 DMO; Maui ......................................................

..................................................... Hylaeus takumiae, new species (p. 195)


52(51). Mandible normal, with two teeth (Fig. 9A) ..................................................... 53— Mandible modified, with three teeth or none (Figs. 9B, 9C) ........................... 54

53(52). Vertex and scutum hair short to medium, 6-12; Hawaii, Maui .................................................................................. Hylaeus psammobius (Perkins) (p. 170)

— Vertex and scutum hair long, 12-20 and 10-18 respectively; Hawaii, Maui ..................................................... Hylaeus coniceps (Blackburn), in part (p. 61)

54(52). Mandible with three similar-sized teeth (Fig. 9B); scutum hair short, 2-6; coast ofHawaii, Maui Nui (including Kahoolawe), Oahu ................................................................................................ Hylaeus anthracinus (F. Smith) (p. 52)

— Mandible straight across at apex, as wide or wider at apex than at articulation(Fig. 9C); scutum hair medium to long, 8-20; mountains of Hawaii ..................................................................... Hylaeus rugulosus (Perkins), (p. 176)

55(50). Upper frons with dense granular sculpture and minute transverse wrinkles,obscuring punctation medially in frontal view, only very slightly lustrous;large species, wing length usually over 6 mm ........................................... 56

— Upper frons ground almost smooth to coriaceous, punctation obvious, morestrongly lustrous; smaller species, wing length less than 6 mm ................ 57

56(55). Maui Nui (including Kahoolawe), Oahu .. Hylaeus assimulans (Perkins) (p. 55)— Hawaii .......................................................... Hylaeus ombrias (Perkins) (p. 158)

57(55). Punctation of upper frons denser medially, with crowded, distorted pits aroundmedian line ................................................................................................ 58

— Punctation of upper frons almost even, pits sometimes close above antennal socketbut not crowded and distorted; Hawaii, Maui, Lanai .................................................................................... Hylaeus flavipes (F. Smith), in part (p. 90)

58(57). Upper frons strongly lustrous, almost shining, especially laterally; rare, Hawaii,Lanai, Molokai .. Hylaeus niloticus (Warncke), new combination (p. 152)

— Upper frons only slightly lustrous, even laterally; Maui Nui, Oahu ............................................................................... Hylaeus longiceps (Perkins) (p. 133)

HAWAIIAN HYLAEUS 43

TAXONOMY OF HYLAEUS (NESOPROSOPIS) SPECIES

Hylaeus (Nesoprosopis) akoko Magnacca and Daly, new species(Fig. 10)

DIAGNOSIS. Large bees with punctured terga. Males with a large ivory mark on thescape, irregular apical spots on the clypeus (probably some individuals will be found witha complete transverse line, or unmarked), and orange and ivory marks on the legs. Scapestrongly arched. Most of body with close, deep punctation. Similar to H. pubescens, differingin the more extensive markings, nearly straight lateral edge of scape, lack of a distincthump on S2, and lack of densely plumose golden or orange hair on T5-T7.

Distribution. Hawaii; only one specimen known, from mesic forest.Male. Measurements: Malar space, 8; eye, 180:134:136:96; clypeus, 96×86; frons,

36×34:108; interalveolar, 24:32:54:62, D 24; interocellar, 30:36:36:42, D 18; scape, 60×46;wing length, 7.1 mm (holotype).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area and frons withdistinct, nearly round pits about 1 pit width apart or less; ground minutely granular, slightlylustrous; no impunctate area above antennal sockets. Lower paraocular depression distinct.Frontoclypeal sulcus nearly straight. Scape in frontal view moderately dilated, width morethan half length, ratio about 0.8; lateral edge in frontal view nearly straight; strongly archedin lateral view with maximum arch about two-thirds along; underside with a narrow groove;with medium-length, pale, simple hair. Vertex hair 16-24, white, simple.

Mesosoma. Scutum with distinct pits 1 pit width apart or less, ground granular, dull;hair medium to long (10-16), white, simple. Scutellum similar, with pits slightly larger.Mesepisternum with pits distinct, round, less than 1 pit width apart, and long, pale brownishwhite, plumose hair; ground granular, dull. Hypoepimeron punctured, slightly lustrous,distinctly swollen. Metanotum dull. Basal area of propodeum slightly protuberant in middle,roughened overall by reticulate rugae; rugae and ground minutely wrinkled, slightly lustrous;lateral carina strong, terminating anterolaterally in a protuberant area.

Metasoma. Disk of T2 with distinct punctation, pits less than 1 pit width apart; withfine, pale, appressed hairs denser, longer, plumose, and white at posterolateral margins;ground smoothly coriaceous, lustrous. Pits deeper, more distinct on T3-7. S2 evenly convex,but without a distinct hump. T7 hair dark reddish brown, prostrate; T5-6 hair not particularlyplumose or dense.

Terminalia. Lateral process of S7 tapered, with length subequal to width of hemisternite.Median process of S8 strongly dilated, apex bifurcate with broad, rounded tips. Apices ofgonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face of holotype with small,obscure ivory spots along apical margin of clypeus, probably varying from unmarked to atransverse stripe; scape in frontal view ivory on lateral third. Pronotum with narrowlyinterrupted ivory transverse collar stripe; lobes unmarked. Legs marked as follows: foreand mid tibiae and femora orange anteriorly (similar to typical pale area), fore and midtibiae with small basal and apical ivory marks; hind tibia with an ivory stripe along entire


dorsal length, narrowly interrupted about three-quarters to apex; all basitarsi ivory; midand hind second tarsal segments with ivory spot. Wings dark smoky.

Female. Unknown.Holotype ♂ labeled “HI: Hawaii Is., Puu Waawaa, 4100', on Chamaesyce olowaluana.

K. Magnacca, 1 August 2002, KM-02-092” (CUIC).Flower records. Chamaesyce olowaluana.Etymology. The species epithet akoko comes from the Hawaiian name for Chamaesyce.Remarks. This appears to be a relative of H. pubescens adapted to drier habitats. Although

only a few C. olowaluana trees have been collected on, they can support a remarkabledensity of bee species; eight species were collected in company with this one, including therare H. dimidiatus, H. filicum, and H. hula. Unfortunately these formerly common treesare not reproducing because seedlings are eaten by sheep.

Fig. 10. Hylaeus akoko, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E.

A B C D

0.5 mm

E F

1.0 mm

HAWAIIAN HYLAEUS 45

Hylaeus (Nesoprosopis) andrenoides (Perkins)(Fig. 11)

Nesoprosopis andrenoides Perkins, 1899:111 (♀; Pl. 2, Fig. 6; flower: Metrosideros).Hylaeus (Nesoprosopis) andrenoides, Meade-Waldo, 1923:35 (list).Nesoprosopis andrenoides, Daly, 1994:116 (lectotype ♀ BMNH 17a 145; Kauai, Waimea,

2000 ft; wing length, 7.6 mm).

DIAGNOSIS. Large bees with punctured terga and dark wings, terga with unique apicalbands of white pubescence, and unique body color with blue reflections. Male face with aparallel-sided stripe and with extremely dilated scape.

Distribution. Kauai; montane mesic and wet forest.Male. Measurements: Malar space, 7; eye, 154:142:140:93; clypeus, 83×81; frons,

25×32:98; interalveolar, 20:34:56:66, D 20; interocellar, 32:42:29:44, D 20; scape, 56×69;wing length, 6.2 mm (Kauai, Alakai Swamp Trail, 3600 ft, KM-99-212).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area, frons, andmiddle uniformly with contiguous, slightly distorted pits; ground largely eliminated by pits,lustrous; frontal line present; small impunctate, lustrous area above antennal socket. Lowerparaocular depression distinct, minutely wrinkled to smooth, shiny. Frontoclypeal sulcusslightly arched. Scape in frontal view heart-shaped, extremely dilated, width greater thanlength, ratio about 1.2; lateral edge in frontal view strongly convex; strongly arched inlateral view with maximum arch near middle of scape; underside of median edge with abroad, shallow pit near the apex; with long, brownish, simple hair. Vertex hair 16-30, palebrown, mostly simple.

Mesosoma. Scutum with distinct pits 1-2 pit widths apart, ground slightly granular andslightly lustrous; hair medium-length (10-14), brown, mostly simple. Scutellum with slightlylarger pits, smoothly coriaceous, lustrous. Mesepisternum with distinct to indistinct, mostlyround pits, about 1 pit width apart or less, with long, brown, weakly plumose hair; groundminutely coriaceous, slightly lustrous. Hypoepimeron punctured, smooth, lustrous, swollen.Metanotum minutely wrinkled, dull. Basal area of propodeum reticulate to areolate rugoseincluding brow, shiny between rugae; below brow granular, dull.

Metasoma. Disk of T2 impunctate, polished and shiny, generally with fine short hairsspaced about one hair length apart; apical terga with minute pits. Terga 1-3 at sides withconspicuous apical bands of appressed, white, thick, plumose hair. T7 hair long, erect,dark brown.

Terminalia. Lateral process of S7 tapered, slender, length greater than width ofhemisternite. Median process of S8 slightly dilated, apex bifurcate with moderately long,straplike, rounded tips. Apices of gonoforceps exceeding penis valves, strongly curvedinward.

Color. Labrum and mandible black. Face with a narrow, longitudinal stripe: clypeus withcentral yellow stripe full length; supraclypeal area below acme with yellow spot that mayor may not be continuous with clypeal stripe; paraocular area and scape unmarked. Pronotumunmarked. Legs dark brown to black except fore tibia and distal fore femur with paleareas. Mesosoma black with faint metallic blue reflections; metasoma black with distinctblue reflections. Wings dark brown with iridescence.


Female. Head. Malar space medium (ratio about 0.5 DMO). Upper paraocular area,frons, and middle uniformly with round or slightly distorted pits about 1 pit width or lessapart; ground not visible in some bees, if visible among pits then with minute parallelwrinkles curved laterad from beneath ocelli to downward along eye, slightly lustrous; frontalline present. Apex of clypeus shallowly emarginate. Mandible weakly three-toothed, thetwo anterior (dorsal) teeth poorly separated. Dorsal end of facial fovea extended to levelof lateral ocellus. Vertex hair 16-30, white, plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with distinct pits 1-2 pitwidths apart, ground granular, dull; hair long (12-24), white, weakly plumose. Scutellumwith slightly larger pits, ground dull to slightly lustrous. Mesepisternum with distinct, roundpits, 1 pit width apart, and medium-length, brown, weakly plumose hair; ground minutely

Fig. 11. Hylaeus andrenoides, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

A B C D

E F

1.0 mm

HAWAIIAN HYLAEUS 47

wrinkled, slightly lustrous. Hypoepimeron with distinct, round pits, 1-2 pit widths apart,ground minutely wrinkled; slightly swollen. Metanotum dull. Basal area of propodeumreticulately rugose over most of the area including brow, shiny between rugae.

Metasoma. Disk of T2 impunctate, polished and shiny, generally with fine very shorthairs more than a hair length apart; apical terga with minute pits. Conspicuous apical bandsof appressed, thick, white, plumose hair on terga 1-4, thin or interrupted at middle of terga1 and 4, complete on terga 2-3. T6 hair long, prostrate to erect, dark brown.

Color. Head and legs dark brown, except fore tibia with or without pale area; thoraxblack; metasoma black with bluish reflections. Face and pronotum unmarked. Wings darkbrown with iridescence.

Perkins’ collections. Kauai: Waimea, 2000 ft (BMNH).Recent collections. Kauai: Alakai Swamp Trail, 3600 ft; Mohihi Trail, 3700 and 4000

ft; Paaiki Valley, 2700 ft, Pihea Trail.Flower records. Chamaesyce atrococca, Cheirodendron, Claoxylon, Metrosideros,

Psychotria mariniana.Remarks. The unusual blue color and abdominal hair bands of this species make it one

of the most striking Nesoprosopis.References. Perkins, 1913:82 (biogeography).

Hylaeus (Nesoprosopis) angustulus (Perkins)(Fig. 12)

Nesoprosopis angustula Perkins, 1899:95-96 (♂♀).Hylaeus (Nesoprosopis) angustulus, Meade-Waldo, 1923:34 (list).Nesoprosopis angustula, Daly, 1994:116 (lectotype ♂ BMNH 17a 150a; Lanai, Mts.

Koele, 2500 ft).

DIAGNOSIS. Medium-sized bees with clear wings, propodeum with weak basal rugae,otherwise smooth to brow. Male has unusual scape with very slight arch, very short malarspace, frons with dull, dense pits but no dark patch of hair, three separate yellow facialmarks and otherwise unmarked. Female with black body, short malar space, pronotal lobesand collar at extreme sides marked with yellow, sometimes with yellow paraocular marks.

Distribution. Maui (new record) and Lanai; montane wet forest.Male. Measurements: Malar space, 3; eye, 99:89:86:57; clypeus, 52×49; frons, 18×22:71;

interalveolar, 14:17:40:48, D 18; interocellar, 22:26:18:26, D 14; scape, 37×18; wing length,4.0 mm (H. angustulus lectotype).

Head. Malar space very short (ratio about 0.2-0.3 DMO). Upper paraocular area withnearly round, distinct pits about 1 pit width apart, ground with parallel wrinkles; frons withpits becoming closer, smaller, distorted, contiguous and aligned with parallel wrinkles; atmiddle pits very dense and dull (at certain angles a transverse band of dark sculpture isevident), but no dark spot with hairs; no frontal line; large lustrous impunctate area aboveantennal sockets. Lower paraocular depression absent. Frontoclypeal sulcus nearly straightor slightly arched. Scape in frontal view slightly dilated, width half length, ratio about 0.5;


lateral edge in frontal view nearly straight; very slightly arched in lateral view with maximumarch near middle of scape; underside of median edge with a broad depression with granularsculpture; with medium-length, brown, simple hair. Vertex hair 12-20, brown, simple.

Mesosoma. Scutum with indistinct minute pits 1-2 pit widths apart, ground granular,dull; hair very short (2-4), dark brown, simple. Scutellum and metanotum with pits larger,more distinct, 2 pit widths apart and with ground granular, slightly lustrous. Mesepisternumwith indistinct, round pits about 1-3 pit widths apart, and sparse, long, pale brown, weaklyplumose hair; ground smoothly coriaceous, lustrous. Hypoepimeron impunctate, smooth,lustrous; slightly swollen. Basal area of propodeum with short, weak, widely spaced, lineaterugae in basal quarter; anterior area rather long, smoothly coriaceous, lustrous, and withpronounced brow.

Fig. 12. Hylaeus angustulus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

A B C D

E F

1.0 mm

HAWAIIAN HYLAEUS 49

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; largely hairless, butwith some fine, pale, appressed hairs spaced more than a hair length apart. T7 hair long,black, erect.

Terminalia. S7 strongly L-shaped: lateral process almost parallel-sided, widest nearapex, length greater than width of hemisternite; hemisternite very short. Median process ofS8 strongly dilated, apex bifurcate with short, broad, rounded tips. Apices of gonoforcepsexceeding penis valves.

Color. Labrum and mandible brown to black. Head and mesosoma black; metasomadark brown. Face with three separate yellow marks: clypeus with irregular subapical spot;lower paraocular area with mark to level of antennal socket; scape unmarked. Pronotumwith or without yellow marks on lobes. Legs dark brown except fore tibia with pale area,sometimes extended onto apex of femur or base of basitarsus. Wings clear.

Female. Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area withdistinct, round pits about 1 pit width apart; on frons pits becoming smaller but still 1 pitwidth apart, ground minutely wrinkled and lustrous; at middle pits contiguous, distorted,separated by longitudinal wrinkles. Frontal line faint. Apex of clypeus shallowly emarginate.Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 10-16, brown,mostly simple.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with extremely small, obscurepits more than 2 pit widths apart, ground granular, dull; hair very short (2-4), brown,simple. Scutellum with pits larger, more distinct, 2 pit widths apart and with ground granular,slightly lustrous. Mesepisternum with indistinct pits 1-3 pit widths apart, and yellowishwhite, weakly plumose hair; ground smooth, lustrous. Hypoepimeron impunctate, smooth,lustrous; slightly swollen. Metanotum with indistinct pits less than 1 pit width apart, groundgranular and dull. Basal area of propodeum with short, weak, widely spaced, lineate rugaein basal quarter; anterior area long, smoothly coriaceous, lustrous, and with pronouncedbrow.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with large bare areas,otherwise with fine, short, pale, appressed hairs spaced more than a hair length apart. T6hair long, dark brown, erect.

Color. Head and mesosoma black; legs and metasoma dark brown. Face with smallparaocular marks or unmarked. Pronotum with lobes yellow and with yellow spots atextreme sides of collar. Legs with fore tibia with obscure pale area, all tibiae with or withoutyellow marks at base. Wings yellowish, nearly clear.

Perkins’ collections. Lanai: Mts. Koele, 2500 ft (BMNH).Recent collections. Maui: Makawao Forest Reserve, 3600-4300 ft.Flower records. Melicope sp., Rubus penetrans*.Remarks. Although this species has not been recollected on Lanai, recent specimens

from East Maui match them closely. It is very close to H. dumetorum from Hawaii, includingthe unique broad groove on the underside of the scape, but lacks the conspicuous patch oflong black hair on the frons of that species.

References. Perkins, 1910:602 (key).


Hylaeus (Nesoprosopis) anomalus (Perkins)(Fig. 13)

Nesoprosopis anomala Perkins, 1899:112-113 (♀; holotype ♀ BMNH 17a 146; Oahu,Honolulu, 2000 ft, wing length, 5.6 mm; flower: Acacia koa).

Hylaeus (Nesoprosopis) anomalus, Meade-Waldo, 1923:35 (list).Nesoprosopis anomala, Daly, 1994:116 (holotype ♀ confirmed).

DIAGNOSIS. Medium-sized bees with clear to smoky wings; scutum with unusual densepunctation less than 1 pit width apart; unique long, pale yellow-brown, plumose hair onmesosoma; may have some red on metasomal terga; and punctured terga. Male with yellowfacial mark only at apex of clypeus or extended at sides in a narrowing stripe above antennalsockets; clypeal mark transverse and apical, or extended dorsally to the supraclypeal area.Face marks similar to H. gliddenae, from which it differs by the mostly black metasomaand the shape of the scape and terminalia. Fresh male specimens are said to have T1-3orange red at base. Female with unique red head, antennae, and T1.

Distribution. Oahu; montane mesic and wet forest. No recent collections.Male. Measurements: Malar space, 7; eye, 141:124:120:84; clypeus, 77×69; frons,

23×31:90; interalveolar, 22:28:51:61, D 22; interocellar, 33:33:26:33, D 17; scape, 46×32;wing length, 5.1 mm (Oahu, Waimea, 1800 ft, BPBM 3221).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area with round,contiguous pits; a broad area of frons above antennal sockets with smaller, contiguous pitsand dark hair creating a large somewhat dark area. Lower paraocular depression absent.Frontoclypeal sulcus slightly arched. Scape in frontal view moderately dilated, width morethan half length, ratio about 0.7; lateral edge in frontal view nearly straight; strongly archedin lateral view with maximum arch near middle of scape; no groove on underside; withmedium-length, pale yellow-brown, mostly weakly plumose hair. Vertex hair 20-30, paleyellow-brown, mixed simple and weakly plumose.

Mesosoma. Scutum with distinct, nearly round pits, less than 1 pit width apart, groundminutely wrinkled and dull; hair medium-length (10-16), pale yellow-brown, plumose.Mesepisternum with round pits less than 1 pit width apart, and long, pale yellow-brown,plumose hair; ground smooth to minutely wrinkled, slightly lustrous. Hypoepimeronpunctate, smooth, slightly lustrous; slightly swollen. Metanotum dull. Basal area ofpropodeum reticulately rugose over entire area; shiny between rugae; sides of propodeumwith dense, long, pale brown, plumose.

Metasoma. Disk of T2 with distinct punctation consisting of minute, shallow pits about2 pit widths apart; smoothly coriaceous, lustrous; with fine, short, pale, appressed hairs.T7 hair medium-length, brown, prostrate.

Terminalia. Lateral process of S7 tapered, with length subequal to width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with broad, rounded tips. Apicesof gonoforceps subequal to penis valves.

Color. Labrum and mandible yellowish brown to brown. Face with only a narrow yellowline at apex of clypeus or with a “U” or trident shape facial mark: clypeus with irregulartransverse yellow mark, often extended basally in a narrow line as far as acme; paraocular

HAWAIIAN HYLAEUS 51

area with yellow band along the eye to well above antennal socket; scape unmarked butyellowish brown. Pronotum with transverse collar stripe complete; lobes unmarked. Legsorange to yellowish brown except hind tibia with obscure yellow stripe in basal third ofsegment. Abdominal terga with or without red on T1-3. Wings clear.

Female. Head. Malar space medium-length (ratio about 0.5 DMO). Upper paraoculararea and frons with contiguous, subpolygonal pits uniformly across areas; a few longitudinalwrinkles at the middle; no frontal line evident. Apex of clypeus shallowly emarginate.Dorsal end of facial fovea extended just to level of lateral ocellus. Vertex hair 20-30, palebrown, plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with distinct, nearly roundpits, less than 1 pit width apart, ground minutely wrinkled, dull; hair medium-length (8-

Fig. 13. Hylaeus anomalus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

F

A B C D

E

1.0 mm


16), plumose, pale yellow-brown. Mesepisternum with round or slightly distorted pits lessthan 1 pit width apart or contiguous, and yellow to brownish white, plumose hair; groundnearly smooth, lustrous. Hypoepimeron punctate, nearly smooth, lustrous; slightly swollen.Metanotum dull. Basal area of propodeum reticulately rugose over entire area; shiny betweenrugae.

Metasoma. Disk of T2 with distinct punctation consisting of minute, shallow pits about2 pit widths apart; smoothly coriaceous, lustrous; with fine, short, pale, appressed hairs.T6 hair long, dark brown, erect.

Color. Head and antennae red; mesosoma black; metasoma with first segment red, theremainder dark reddish black. Face unmarked. Pronotum with ivory transverse collar stripecomplete; lobes unmarked and black. Tegula with ivory spot. Legs red except: fore tibiawith basal ivory spot; mid tibia with two-thirds ivory stripe; and hind tibiae with ivorystripe almost full length. Wings smoky.

Perkins’ collections. Oahu: Honolulu, 2000 ft (BMNH); Tantalus (BPBM, OUMNH);Waimea, 1800 ft (BPBM).

Recent collections. None.Flower records. Acacia, Psychotria sp.Remarks. Colors of the integument and pubescence are based on old specimens. Fullaway

(1918) describes the male with orange red legs and basal abdominal terga, but specimenswe have seen have the legs yellowish brown and the terga reddish brown or without adistinct red color on the abdomen. The marks of the male are yellow; those of the femaleare ivory. Although most species with punctured terga have the scutum punctation relativelyclose, this species has the most extreme development of this character.

References. Perkins, 1907a:43 (record: Oahu, Tantalus); 1910:604 (key); 1913:82(biogeography). Fullaway, 1918:397-398 (♂♀). Swezey, 1922:14 (nest: in Pipturus tree;odor of citronella; flower: Psychotria (referred to as Straussia); record: Oahu, Tantalus);1954:39, 167 (nest: in twigs of Boehmeria grandis and Pipturus; record: Oahu, Tantalus).Meade-Waldo, 1923:35 (list). Williams, 1927:427 (nest: in dead twigs of Pipturus andBoehmeria; flower: Acacia koa).

Hylaeus (Nesoprosopis) anthracinus (F. Smith)(Fig. 14)

Prosopis anthracina F. Smith, 1853:23 (♂♀).Nesoprosopis anthracina, Perkins, 1899:100-101 (♂♀; synonymy; Pl. 2, Fig. 17).Hylaeus (Nesoprosopis) anthracinus, Meade-Waldo, 1923:34 (list, synonymy).Prosopis anthracina, Daly, 1994:119 (lectotype ♀ BMNH 17a 172; Sandwich Islands;

wing length, 4.1 mm).Prosopis rugiventris Blackburn, 1886:146-148 (♂♀; key) (synonymy, Perkins 1899:100).Prosopis rugiventris, Daly, 1994:113 (types not located).

DIAGNOSIS. Medium-sized bees with clear to smoky wings, black legs. Male face withoval yellow mark or lower face all yellow, not reaching above antennal sockets; scape

HAWAIIAN HYLAEUS 53

Fig. 14. Hylaeus anthracinus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

A B C D

E F

1.0 mm

moderately to strongly dilated. Female is entirely black. Usually occurs in company withH. flavipes or H. longiceps, from which the females can be distinguished by the black hairson the apex of the abdomen and the unusual three-toothed mandible, a feature shared onlywith H. flavifrons. Closely related to H. flavifrons, from which the male differs in the lessdilated scape, undilated process of S8, and usually smaller facial markings.

Distribution. Hawaii, Maui, Kahoolawe, Lanai, Molokai, and Oahu; coast and drylowlands. No recent collections from Lanai.

Male. Measurements: Malar space, 5; eye, 148:113:110:77; clypeus, 77×74; frons,28×26:95; interalveolar, 18:28:51:61, D 20; interocellar, 31:28:18:26, D 15; scape, 46×34;wing length, 4.3 mm, tip damaged (Maui, BPBM 3224).


Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area and frons ratherflat and uniformly covered with round, shallow pits about 1 pit width apart, interrupted atthe middle in some bees by faint, incomplete frontal line; impunctate area above antennalsockets quite small, pits evident close to socket; ground minutely wrinkled, but smooth,lustrous. Lower paraocular depression distinct, but shallow. Frontoclypeal sulcus slightlyarched. Scape in frontal view moderately to strongly dilated, width more than half length,ratio about 0.7-0.9; in moderately dilated scapes lateral edge in frontal view nearly straightin middle and rounded only at ends, and in strongly dilated scapes lateral edge distinctlyrounded (convex); moderately arched in lateral view with maximum arch near middle ofscape; no groove on underside; with short, white, weakly plumose hair. Vertex hair 6-10,white, plumose.

Mesosoma. Scutum with distinct pits about 1-2 pit widths apart, ground of scutum andscutellum smoothly coriaceous, lustrous; hair short (4-6), white to pale brown, weaklyplumose. Mesepisternum with distinct, shallow, round pits, about 1 pit width apart, andshort, yellowish white, weakly plumose hair; ground minutely coriaceous, slightly lustrous.Hypoepimeron with few obscure pits, lustrous, slightly swollen. Metanotum minutelywrinkled, dull. Basal area of propodeum lineately rugose or somewhat reticulate to brow,shiny between rugae; ground below brow slightly granularly coriaceous, slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, short, pale,appressed hairs spaced about one hair length apart. T7 hairs dark brown, medium-length,erect.

Terminalia. Lateral process of S7 tapered, broad, length much greater than width ofhemisternite. Median process of S8 linear or slightly dilated just distal to flexure, apexbifurcate with narrow tips longer than broad. Apices of gonoforceps subequal to penisvalves.

Color. Labrum and mandible blackish brown to black. Face with a single large spot:clypeus entirely yellow or nearly so; lower paraocular area with a stripe along sulci ofvariable width, usually relatively broad; supraclypeal area to acme yellow; scape unmarked.Pronotum unmarked. Legs dark brown to black, unmarked except pale area on fore tibia.Wings nearly clear to smoky.

Female. Head. Malar space medium-length (ratio about 0.5 DMO). Upper paraoculararea and frons rather uniformly with round, shallow pits about 1 pit width apart, groundminutely wrinkled, slightly lustrous; fine frontal line may be evident. Apex of clypeusshallowly emarginate. Apex of mandible straight, with three small but usually distinct teeth(see Fig. 9B). Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 4-8, white, plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with distinct pits about 1-2pit widths apart, ground of scutum and scutellum smoothly coriaceous, lustrous; hair veryshort (2-6), white to pale reddish brown, mostly simple. Mesepisternum with distinct,shallow, round pits, about 1 pit width apart, and medium-length, yellowish white, weaklyplumose hair; ground minutely coriaceous, slightly lustrous. Hypoepimeron with few obscurepits, lustrous, slightly swollen. Metanotum minutely wrinkled, dull. Basal area of propodeumlineately rugose or somewhat reticulate to brow, shiny between rugae; ground below browslightly granularly coriaceous, slightly lustrous to dull.

HAWAIIAN HYLAEUS 55

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, short, pale,appressed hairs spaced about one hair length apart. T6 hair long and erect, dark brown.

Color. Head and mesosoma black; legs and metasoma dark brown; unmarked exceptwith or without pale area on fore tibia. Wings nearly clear to smoky.

Perkins’ collections. Hawaii: Kealakeakua Bay (OUMNH); Kona (BMNH, BPBM).Maui: Kaulawai (BPBM); “Maui” (BMNH, BPBM); Maui coast (OUMNH); Wailuku,sand hills (BPBM). Lanai: Manele (BPBM, OUMNH). Molokai: lower slopes, mtns(OUMNH); plains (BMNH, BPBM, OUMNH). Oahu: Honolulu (BPBM), coast (BPBM),mts. (BPBM); Waialua (BMNH); coast (BPBM); Waianae (BMNH, BPBM); Waikiki(BPBM, OUMNH).

Recent collections. Hawaii: South Point, coast. Maui: Kanaio Natural Area Reserve,2000 ft; Manawainui West, coast. Kahoolawe: Pali o Kalapakea, 1000 ft. Molokai:Moomomi, coast. Oahu: Kaena Point Natural Area Reserve, coast.

Flower records. Argemone glauca (Maui), Chamaesyce celastroides, C. degeneri,Myoporum, Scaevola sericea, Sesbania tomentosa, Sida, Tournefortia*.

Remarks. This once-widespread species appears to have declined precipitously with theloss of coastal habitat, and now appears to be restricted to one or a few localities on eachisland. However, it appears to have a significant population on the small, uninhabited islandof Kahoolawe. Hylaeus anthracinus is not closely related to the other species (e.g., H.difficilis) with an undilated S8 process and a single central facial mark. The female hasthree small teeth on the mandible, rather than two large, a unique feature shared only withits sister species, H. flavifrons (sometimes other species may appear similar due to wear).It typically occurs in company with H. longiceps or H. flavipes, from which the femalescan also be distinguished by the erect dark hairs of T6.

References. Blackburn and Cameron, 1886:149 (♂♀); 1887:208-209 (♂♀). Dalla Torre,1896:14 (list). Perkins, 1899:104 (probable host of cleptoparasite: N. hilaris); 1907a:43(record: Oahu, Tantalus); 1910:602 (key); 1913:64, 81 (biology, biogeography,morphology). Bridwell, 1919:148 (morphology). Beardsley and Tuthill, 1959:60 (recordof H. anthracinus from Niihau is probably H. flavifrons; see remarks under H. flavifrons).Suehiro, 1986:41, 60 (record: Hawaii, Kona). Hopper and Daly, 1995:35 (record of H.anthracinus from Kauai is H. solaris). Prosopis rugiventris: Blackburn, 1887:206-208(♂♀; key). Perkins, 1899:100 (synonym of N. anthracina). Meade-Waldo, 1923:34 (insynonymy).

Hylaeus (Nesoprosopis) assimulans (Perkins)(Fig. 15)

Nesoprosopis assimulans Perkins, 1899:101-102 (♂♀; Pl. 2, Fig. 19).Hylaeus (Nesoprosopis) assimulans, Meade-Waldo, 1923:34 (list).Nesoprosopis assimulans, Daly, 1994:116 (lectotype ♂ BMNH 17a 165a; Lanai, Mts.

Koele, 2000 ft, wing length, 5.6 mm).Nesoprosopis assimulans oahuensis Perkins, 1899:102 (♂♀) (new synonymy).Hylaeus (Nesoprosopis) assimulans oahuensis, Meade-Waldo, 1923:34 (list).Nesoprosopis assimulans oahuensis, Daly, 1994:116 (lectotype ♂ BMNH 17a 166b;

Oahu, Waianae; wing length, 5.4 mm).


DIAGNOSIS. Medium to large bees with slightly smoky to smoky wings and short malarspaces. Male with unusual scape, slightly dilated, widest near the middle and smaller atboth ends, and with short plumose hair along the median edge; distinguished from larger,more melanic sister species, H. ombrias, by larger facial marks with clypeus always markedand sometimes also the supraclypeal area, lineate rugosity mainly at front half of propodeum,and smoky, but not dark, wings. Female black and unmarked, closely resembling H. longicepsand H. flavipes; differing in sculpture of the frons (see key), and usually much larger.

Distribution. Maui, Lanai, and Oahu; coast and dry lowlands. No recent collectionsfrom Oahu.

Male. Measurements: Malar space, 8; eye, 157:118:112:86; clypeus, 78×82; frons,29×27:106; interalveolar, 16:25:55:67, D 24; interocellar, 35:31:24:31, D 18; scape, 55×27;wing length, 5.6 mm (H. assimulans lectotype).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area with indistinct,round pits about 1 pit width apart, some bees with very weak, parallel wrinkles curvedfrom near lateral ocellus to along the eye; frons with pits becoming more distinct, smaller,and closer medially; middle area with pits contiguous, some pits distorted; central groundnearly smooth, slightly lustrous; no frontal line or other special sculpture at middle;impunctate, lustrous area above antennal sockets. Lower paraocular depression absent.Frontoclypeal sulcus straight or nearly so; supraclypeal area variable in shape, W/L ratio0.8-1.0. Scape in frontal view slightly dilated, widest near the middle, width half of thelength, ratio about 0.5; lateral edge in frontal view nearly straight; slightly arched in lateralview with maximum arch near the middle; no groove on underside; with short, yellowishwhite, plumose hair. Vertex hair 10-14, brown, plumose.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground nearly smooth, slightly lustrous;hair short (4-6), yellowish white, plumose. Mesepisternum with shallow, indistinct pitsabout 1 pit width apart, and long, white, plumose hair; ground smoothly coriaceous, lustrous.Hypoepimeron impunctate, smooth, lustrous; slightly swollen. Metanotum dull. Basal areaof propodeum with weak, sparse, irregular, lineate rugae halfway or more to brow, shinybetween rugae, faintly reticulate behind; brow nearly smoothly coriaceous or minutelywrinkled, lustrous or slightly so.

Metasoma. Disk of T2 impunctate, smoothly coriaceous; with abundant, fine, short,pale, appressed hairs spaced about one hair length apart. Hairs of T7 pale brown, short,appressed.

Terminalia. Lateral process of S7 parallel-sided, length greater than width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with medium-length, narrow,straplike tips. Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face entirely yellow below theantennae, or the clypeus with narrow to broad black margins making three separate spots;the paraocular marks extended at sides in narrowing stripe to above or just below antennalsockets. Supraclypeal area sometimes with a yellow mark; scape unmarked. Pronotumunmarked. Legs dark brown to black except fore tibia with pale area and with or withoutobscure basal yellow spots on all tibiae. Wings slightly smoky to smoky.

HAWAIIAN HYLAEUS 57

Female. Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area andfrons with indistinct, round pits about 1 pit width apart across the areas; ground at sideswith very weak, parallel wrinkles curved from near lateral ocellus to along the eye; centralground of frons medially with weak, transverse, parallel wrinkles; faint frontal line evidentin some bees. Apex of clypeus shallowly emarginate. Dorsal end of facial fovea extendedto level of lateral ocellus. Vertex hair 6-8, pale brown, plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with indistinct, minute, roundpits about 2 or more pit widths apart, ground granular to smooth, dull to lustrous in somebees; hair short (4-6), white to pale brown, plumose. Mesepisternum with shallow, indistinctpits about 2 pit widths apart, and yellowish white, plumose hair; ground striate to nearlysmooth, slightly lustrous. Hypoepimeron impunctate, smooth to weakly striate, slightly

Fig. 15. Hylaeus assimulans, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

A B C D

E F

1.0 mm


lustrous; slightly swollen. Metanotum dull. Basal area of propodeum with irregular,somewhat sparse, lineate rugae halfway or more to brow; surface shiny between rugae;brow nearly smoothly coriaceous or minutely wrinkled, lustrous or slightly so.

Metasoma. Disk of T2 impunctate, smoothly coriaceous; with fine, very short, pale,appressed hairs spaced more than a hair length apart. T6 hair light yellowish brown, medium-length, appressed.

Color. Head and mesosoma dark brown to black; legs and metasoma dark brown;unmarked except fore tibia with pale area. Wings smoky.

Perkins’ collections. Hawaii: Kona coast (BPBM). Maui: “Maui” (BPBM); Wailuku(BMNH, BPBM). Lanai: Awalua (BMNH, BPBM); Mts. Koele, 2000 ft (BMNH). Oahu:Kaala Mts., 500 ft (BPBM); “Oahu” (OUMNH); Waianae (BMNH, BPBM); w. coast, nrsea level (OUMNH); Waianae Mts (BMNH, BPBM).

Recent collections. Maui: Lahainaluna, 1800 ft; Waikapu, Kaohonua, 120 m.Kahoolawe: Kamohio, coast. Lanai: Polihua Rd., 1000 ft; Manele Rd., 600 ft.

Flower records. Lipochaeta lobata, Sida.Remarks. Perkins characterized the populations on Oahu as having the males with yellow

face marks often much reduced and the wings generally darker; the females with wingsentirely infuscate and iridescent in certain lights. He described this type as H. assimulansoahuensis.

Male H. assimulans and the sister species, H. ombrias, approximately resemble H.niloticus in color and overlap in body size. The sister species both have the greatest widthof the scape near the middle; short, plumose hair along the median edge of the scape; andthe median area immediately above the antennal sockets sparsely punctured to impunctate.Hylaeus niloticus has the greatest width of the scape near the apex, mostly simple hairalong the median edge, and the median area above the antennal sockets densely punctured.Although H. niloticus is usually much smaller, small specimens of the others have beencaught.

References. Perkins, 1899:104 (host of cleptoparasite: N. hilaris); 1910:604 (key);1913:81 (host of N. hilaris). Giffard, 1908:180 (record: Lanai). Fullaway, 1918:394(morphology). Suehiro, 1986:60 (record: Hawaii, Kona). Hylaeus assimulans oahuensis:Perkins, 1899:105 (host of cleptoparasite: N. volatilis); 1910:604 (key); 1913:81 (host ofcleptoparasite: N. volatilis). Meade-Waldo, 1923:34 (list). Fullaway and Krauss, 1945:169-170 (biology of subgenus).

Hylaeus (Nesoprosopis) chlorostictus (Perkins)(Fig. 16)

Nesoprosopis chlorosticta Perkins, 1899:78-79 (♂♀).Hylaeus (Nesoprosopis) chlorostictus, Meade-Waldo, 1923:33 (list).Nesoprosopis chlorosticta, Daly, 1994:116 (lectotype ♂ BMNH 17a 156b; Kauai,

Waimea, 2000 ft; wing length, 4.4 mm).

DIAGNOSIS. Medium-sized bees with smoky wings and short malar spaces. Male facewith oval yellow mark divided by fine black sulci, legs marked with yellow, unusual lateral

HAWAIIAN HYLAEUS 59

process of S7 long and broad, sword-shaped, median process of S8 not dilated and withlong narrow tips, unusual gonoforceps expanded apically beyond the penis valves. Femaleblack and unmarked, distinguishable from all others by the unique line down the middle ofthe clypeus. Closely related to H. facilis and H. simplex.

Distribution. Kauai; coast, lowland and montane dry shrubland, dry and mesic forest.Male. Measurements: Malar space, 6; eye, 141:120:111:76; clypeus, 69×69; frons,

31×28:95; interalveolar, 12:29:51:61, D 22; interocellar, 31:31:26:31, D 17; scape, 57×28;wing length, 4.7 mm (Kauai, Waimea, 2000 ft, BPBM 3286).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area and frons withindistinct, round, shallow pits less than 1 pit width apart; ground minutely wrinkled, dull;median, impunctate, lustrous area just above antennal sockets; frontal line indistinct. Lowerparaocular depression distinct, but shallow. Frontoclypeal sulcus arched. Scape in frontalview slightly dilated, width half length, ratio about 0.5; lateral edge in frontal view nearlystraight or slightly concave; strongly arched in lateral view with maximum arch near middleof scape; no groove on underside; with medium-length, brownish white, weakly plumosehair along the median edge. Vertex hair 14-20, brown, plumose.

Mesosoma. Scutum with pits about 2 pit widths apart, ground minutely granularlycoriaceous, dull; hair medium-length (8-12), pale brown, plumose. Scutellum with pitsslightly larger, ground smoothly coriaceous, slightly lustrous. Mesepisternum with indistinct,shallow, round pits, about 1 pit width apart, and long, white, plumose hair; ground minutelycoriaceous, slightly lustrous. Hypoepimeron impunctate, lustrous, nearly flat. Metanotumdull. Basal area of propodeum lineately rugose to brow, shiny between rugae at front;brow granularly coriaceous, dull to slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, pale,appressed hairs spaced about one hair length apart. T7 hairs short, dark brown, prostrate.

Terminalia. Lateral process of S7 broad, sword-shaped, length much greater than widthof hemisternite. Median process of S8 linear, apex bifurcate with long narrow tips. Apicesof gonoforceps exceeding penis valves.

Color. Labrum and mandible blackish brown to black. Face with a single large spot:clypeus entirely yellow or nearly so; lower paraocular area with a narrow yellow stripealong sulci below antennal socket; supraclypeal area to acme yellow; scape unmarked.Pronotum unmarked. Legs dark brown to black except: fore tibia with pale area, and withor without a trace of yellow at base; mid tibia variable, with or without a trace of yellow atbase; hind tibia with basal yellow stripe less than half length; mid and hind basitarsi variable,with or without some yellow marking at base. Wings smoky.

Female. Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area andfrons with round, shallow pits about 1 pit width apart or less, slightly smaller and closertoward the middle; ground minutely wrinkled, slightly lustrous; fine frontal line evident.Apex of clypeus distinctly emarginate. Clypeus with a faint longitudinal line down middleconsisting of a depression or narrow impunctate area, most apparent near frontoclypealsulcus and usually visible at low angles. Dorsal end of facial fovea extended to level oflateral ocellus. Vertex hair 14-24, brown, plumose.


Mesosoma. Fore tarsus with hairs apically curved. Scutum with pits about 2 pit widthsapart, ground minutely granularly coriaceous, dull; hair medium-length (8-12), pale brown,plumose. Scutellum with pits slightly larger, ground smoothly coriaceous, slightly lustrous.Mesepisternum with indistinct, shallow, round pits, about 1 pit width apart, and long,white, plumose hair; ground minutely coriaceous, slightly lustrous. Hypoepimeronimpunctate, lustrous, nearly flat. Metanotum dull. Basal area of propodeum lineately rugoseto brow, shiny between rugae at front; brow smoothly coriaceous, lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, pale,appressed hairs spaced about one hair length apart. T6 hair short and dense, dark brown,prostrate.

Fig. 16. Hylaeus chlorostictus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

A B C D

E F

1.0 mm

HAWAIIAN HYLAEUS 61

Color. Head and mesosoma dark brown to black; legs and metasoma dark brown;unmarked. Wings smoky.

Perkins’ collections. Kauai: Halemanu, 4000 ft (BPBM); “Kauai,” 1500 ft (OUMNH);Lihue, nr (BPBM);Waimea, 2000 ft (BMNH, BPBM).

Recent collections. Kauai: Kokee Rd., 3600 ft; Paaiki Valley, 2700 ft; Milolii Rd.,2700 ft; Kukui Trail, 2600 ft; Polihale SP, coast.

Flower records. Acacia, Claoxylon, Myoporum, Styphelia.Remarks. The measured male of H. chlorostictus has extreme ratios of “socket diameter/

interantennal distance” at 1.9 and “antenna-eye distance/interantennal distance” at 2.6,indicating that the antennal sockets are extremely close together. The length ratio clypeus/subantennal sulcus at 2.3 is also unusually high.

References. Perkins, 1910:601 (key); 1913:64 (biogeography).

Hylaeus (Nesoprosopis) coniceps (Blackburn)(Fig. 17)

Prosopis coniceps Blackburn, 1886:145-146 (♂♀; holotype ♂ BPBM 2339; Hawaii,Mauna Kea, 7000 ft; wing length, 4.3 mm).

Nesoprosopis coniceps, Perkins, 1899:91-92 (♂♀).Hylaeus (Nesoprosopis) coniceps, Meade-Waldo, 1923:34 (list).Nesoprosopis coniceps, Daly, 1994:113 (holotype ♂ confirmed).Nesoprosopis comes Perkins, 1899:90-91 (♂♀) (new synonymy).Hylaeus (Nesoprosopis) comes, Meade-Waldo, 1923:34 (list).Nesoprosopis comes, Daly, 1994:116 (lectotype ♂ BMNH 17a 161a; Maui, Haleakala,


DIAGNOSIS. Medium-sized bees with clear to slightly smoky wings. Male usually withyellow marks on the clypeus and sometimes lower paraocular areas (most Hawaii specimenswith the latter). Distinct from other three-spot species by the combination of long scutumhair and lineately rugose propodeum; also usually with paraocular marks smaller than clypeal.Female black and unmarked; resembles H. difficilis, but with longer erect hairs on T6.

Distribution. Hawaii and Maui; widespread from montane mesic and wet forest tosubalpine shrubland.

Male. Measurements: Malar space, 5; eye, 108:108:102:69; clypeus, 59×61; frons,15×26:74; interalveolar, 18:26:44:54, D 17; interocellar, 26 33:18:33, D 13; scape, 39×24;wing length, 4.2 mm (Hawaii, Kau, 4000 ft, BPBM 3295).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area with pits lessthan 1 pit width apart and ground with fine wrinkles along eye, medially on frons with pitscloser, crowded and distorted in subpolygonal shapes; a large, dark, rhomboid spot at themiddle caused by dense black hairs and very dense punctation; shiny area just below darkspot and above antennal socket. Lower paraocular depression very shallow. Frontoclypealsulcus slightly arched. Scape in frontal view slightly to moderately dilated, width morethan half length, ratio about 0.6-0.7; lateral edge in frontal view nearly straight; moderately


arched in lateral view with maximum arch near middle of scape; underside of median edgewith a groove; with long, pale brownish, simple hairs. Vertex hair 12-20, brown, simple.

Mesosoma. Scutum with pits 1-2 or more pit widths apart, ground dull; hair long (16-18), brown, weakly plumose. Scutellum with pits 1-2 or more pit widths apart, lustrous.Mesepisternum with indistinct, shallow, round pits, about 1-2 pit widths apart, and long,brownish white, plumose hair; ground minutely coriaceous, slightly lustrous. Hypoepimeronimpunctate, slightly lustrous, nearly flat. Metanotum dull. Basal area of propodeum withrather long anterior area; weakly lineate or irregular rugose halfway or more to brow,spaced 2-4 ruga widths apart; brow granularly coriaceous and slightly lustrous or dull.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart. T7 hair medium-length, brown, erect.

Fig. 17. Hylaeus coniceps, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.

0.5 mm

A B C D

FE

1.0 mm

HAWAIIAN HYLAEUS 63

Terminalia. Lateral process of S7 tapered, length subequal to width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with broad tips. Apices ofgonoforceps exceeding penis valves.

Color. Head and mesosoma black, metasoma dark brown to black. Labrum and mandibleblackish brown to black. Face with one or three separate marks: clypeus with irregular,subapical yellow spot; lower paraocular area with yellow mark of varying size below antennalsocket (if reduced, present only near apex of clypeus); Hawaii Island specimens almostalways with three spots, Maui specimens usually with one spot or none. Supraclypeal areaand scape unmarked. Pronotum unmarked. Legs dark brown to black except fore tibiawith pale area, basitarsi with or without yellow marking, mid and hind terminal tarsalsegments with or without yellow marking, hind tibia with or without obscure yellow spotat very base. Wings clear to slightly smoky.

Female. Head. Malar space short to medium-length (ratio about 0.4-0.5 DMO). Upperparaocular area and frons convex; upper paraocular area with nearly round pits less than 1pit width apart, ground lustrous to shiny; frons with pits becoming crowded medially,contiguous and partly distorted, and at the middle a few, fine, longitudinal carinulae; frontalline, if any, distinct only near acme; small shiny area just above antennal socket. Apex ofclypeus shallowly emarginate. Dorsal end of facial fovea extended just to level of lateralocellus. Vertex hair 12-20, brown, simple.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with distinct pits 2 or morepit widths apart, ground slightly lustrous; hair medium to long (mostly 10-12 with scatteredhairs up to 18), brown, simple. Scutellum with slightly larger pits, lustrous. Scutum andscutellum appear slightly flattened. Mesepisternum with indistinct, shallow, round pits,about 1-3 pit widths apart, and short, yellow-brownish white, weakly plumose hair; groundminutely coriaceous, slightly lustrous. Hypoepimeron impunctate, slightly lustrous, nearlyflat. Metanotum slightly lustrous. Basal area of propodeum dull to slightly lustrous withindistinct, weakly lineate rugae about halfway to brow, spaced 2-4 ruga widths apart; browsmooth to granularly coriaceous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart. T6 hair medium to long, dark brown,erect.

Color. Head and mesosoma black; metasoma dark brown to black; face, pronotum, andlegs unmarked. Wings clear to slightly smoky.

Perkins’ collections. Nesoprosopis coniceps: Hawaii: Kau (BPBM), 4000 ft (BMNH,BPBM); Kilauea (BPBM), 4000 ft (OUMNH); Olaa (BPBM). N. comes: Maui: Haleakala,5000 ft (BMNH).

Recent collections. Hawaii: Kipuka Puaulu, 4000 ft; Earthquake Tr., 3800 ft; KonaForest Unit of Hakalau National Wildlife Refuge camp, 5300 ft; Puu Waawaa, 4100 and5500 ft; Saddle Road, 5400 ft; Puu Huluhulu (N. Hilo), 6700 ft. Kipuka Maunaiu, 1890 m.Maui: Lauulu Tr., 2010 m; Haleakala N. P. Service Area, 2073 m; Koolau Gap, 6200 ft;Puu Keokea, 7100 ft.

Flower records. Chamaesyce olowaluana, Crocosmia × crocosmiiflora*, Geraniumcuneatum, Metrosideros, Myoporum, Santalum paniculatum, Scaevola chamissoniana,Sophora.


Remarks. Fullaway (1918:394-395) pointed out the possible synonymy of H. conicepsfrom Hawaii with H. comes from Maui. They are here considered as one species under thesenior name, H. coniceps. As noted by Perkins (1899:92), bees from both islands are similarin structure. Perkins’ key (1910:603) and his subsequent support of the distinction of thetwo species (1911:724) depended on the number of facial marks. Although the majority ofbees on the two islands are as described by Perkins, bees have been taken recently on bothislands that have only a clypeal mark (as described for H. comes) or a clypeal mark plusparaocular marks (as described for H. coniceps), or no marks at all. The small number ofbees available for study from each island also exhibits overlapping variation in leg markingand propodeal sculpture. This is the only one of the three-spot species in which the paraocularspots are shorter than the clypeal spot (occasional particularly xanthic individuals mayhave the paraocular spots large).

See remarks for H. dumetorum and female H. difficilis.References. Blackburn and Cameron, 1886:148 (key). Blackburn, 1887:205-206, 208

(♂♀; key). Dalla Torre, 1896:20 (list). Perkins, 1907b:97 (record: Hawaii, Kilauea); Perkins,1910:603 (key); 1911:724 (variation). Fullaway, 1918:394-395 (♂♀; record: Hawaii,Kilauea). Meade-Waldo, 1923:34 (list). Suehiro, 1986:41, 60 (record: Hawaii, Kona).Hylaeus comes: Perkins, 1910:603 (key). Meade-Waldo, 1923:34 (list). Swezey andWilliams, 1932:181 (record: Hawaii, Nauhi Gulch).

Hylaeus (Nesoprosopis) connectens (Perkins)(Fig. 18)

Nesoprosopis connectens Perkins, 1899:85-86 (♂).Hylaeus (Nesoprosopis) connectens, Meade-Waldo, 1923:34 (list).Nesoprosopis connectens, Daly, 1994:116 (lectotype ♂ BMNH 17a 121; Maui, West

Maui Mts., Iao Valley).Nesoprosopis koae Perkins, 1899:85 (♀) (new synonymy).Hylaeus (Nesoprosopis) koae, Meade-Waldo, 1923:34 (list).Nesoprosopis koae, Daly, 1994:117 (lectotype ♀ BMNH 17a 120; Oahu, Honolulu;

wing length, 4.7 mm).Nesoprosopis vicina Perkins, 1899:84-85 (♀) (new synonymy).Hylaeus (Nesoprosopis) insulae Meade-Waldo, 1923:34 (list; replacement name for

Nesoprosopis vicina Perkins, a junior homonym of Hylaeus vicinus Sichel) (newsynonymy).

Prosopis trigona Warncke, 1970:748 (unwarranted replacement name for Nesoprosopisvicina Perkins) (new synonymy)

Nesoprosopis vicina, Daly, 1994:119 (lectotype ♀ BMNH 17a 118; Hawaii, Puna; winglength, 4.6 mm).

DIAGNOSIS. Medium-sized bees with clear to smoky wings, unusual punctation of upperface with pits spaced about 1 pit width apart across face even at middle, supraclypeal areawider than long, basal area of propodeum lineately rugose in basal half and otherwisesmooth to brow. Male with a single central facial mark, usually covering all or most of the

HAWAIIAN HYLAEUS 65

Fig. 18. Hylaeus connectens, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

A B C

E F

1.0 mm

D

clypeus and supraclypeal area, sometimes including adjacent paraocular areas or reducedto only the middle of the clypeus. Females black with pronotal lobes marked or not, similarto H. coniceps, but scutum hair shorter.

Distribution. Hawaii, Maui, Lanai, Molokai (new record), Oahu, and Kauai (new record);montane mesic to wet forest, dry shrubland, coast (Kauai). No recent collections fromMolokai.

Male. Measurements: Malar space, 5; eye, 117:111:102:69; clypeus, 62×62; frons,18×29:80; interalveolar, 18:22:46:55, D 22; interocellar, 28:31:18:31, D 15; scape, 40×26;wing length, 4.3 mm (H. connectens lectotype).

Head. Malar space short (ratio about 0.3-0.4 DMO). Upper paraocular area and fronswith mostly round, shallow pits about 1 pit width apart even near the middle, but pits


variable in size and shape, ground varying from minutely wrinkled and quite dull to smoothand lustrous between pits; a fine frontal line between median ocellus and acme that may bepartly obscured by punctation or fine longitudinal wrinkles. Lower paraocular depressionabsent. Frontoclypeal sulcus straight or slightly arched. Supraclypeal area wider than long,W/L ratio 1.2-1.6. Impunctate area above antennal sockets very small or absent. Scape infrontal view slightly to moderately dilated, width more than half length, ratio about 0.6-0.7, widest subapically; lateral edge in frontal view nearly straight to concave; moderatelyarched in lateral view with maximum arch nearer to apex of scape; no groove on underside;and with medium-length, pale, weakly plumose hairs along median surface. Vertex hair 14-20, pale brown, weakly plumose.

Mesosoma. Scutum with distinct pits, about 1-2 pit widths apart, ground slightly lustrous;hair medium-length (6-12), reddish brown, plumose. Scutellum with pits slightly larger,lustrous. Mesepisternum with distinct, shallow round pits about 1 pit width apart, andyellowish white, plumose hair; ground smooth. Hypoepimeron impunctate, smooth, lustrous,slightly swollen. Metanotum dull or slightly lustrous. Basal area of propodeum lineatelyrugose to basal third or almost to brow, rugae spaced 2-6 ruga widths apart; brow smoothlycoriaceous or slightly granular and slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart. T7 hair medium to long, black, erect.

Terminalia. Lateral process of S7 tapered, pale, length subequal to width of hemisternite.Median process of S8 straight along dorsal edge and without a flexure (one male fromLanai has a distinct arch; some others have a slight arch), moderately dilated, apex bifurcatewith short to moderately long rounded tips. Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face with a single large spot:clypeus entirely yellow or largely so, or with broad black margins; lower paraocular areaunmarked or with yellow spot of variable size and shape adjacent to the clypeus; supraclypealarea unmarked or yellow up to acme. Scape unmarked. Pronotum usually unmarked, somespecimens (especially from Kauai) with yellow on the pronotal lobes. Legs dark brownexcept pale area of fore tibia, and with or without the following marks: apex of fore femur,base of fore and mid tibiae, basal third of hind tibia, and mid and hind basitarsi; front tarsussometimes yellow or light brown. Wings clear to smoky.

Female. Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area andfrons with round, shallow pits about 1 pit width apart, mostly round pits even near themiddle, ground minutely wrinkled and dull varying to smooth and lustrous between pits; afine frontal line between median ocellus and acme that may be obscured by punctation andfine longitudinal wrinkles. Apex of clypeus nearly straight or shallowly emarginate. Dorsalend of facial fovea extended to level of lateral ocellus. Vertex hair 10-16, brown, plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with distinct pits about 2 pitwidths apart, ground lustrous; hair variable, short to medium (4-12), reddish brown, simple.Scutellum with pits slightly larger and farther apart, lustrous. Mesepisternum with shallowpits, sometimes indistinct, about 1-2 pit widths apart, and yellowish white, plumose hair;ground smooth. Hypoepimeron sparsely punctured, nearly flat. Metanotum slightly lustrous.Basal area of propodeum lineately rugose halfway or more to brow, rugae spaced 2-6 rugawidths apart; brow smoothly coriaceous and lustrous.

HAWAIIAN HYLAEUS 67

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart. T6 hair dark brown, long to medium,erect.

Color. Body dark brown to black; unmarked. Legs brown to black; unmarked exceptwith or without pale fore tibia and with or without obscure yellow spot at bases of tibiae.Kauai specimens sometimes with yellow on the pronotal lobes and basal third of hind tibia.Wings clear to smoky.

Perkins’ collections. Nesoprosopis connectens, N. koae, and N. vicina: Hawaii:Hamakua, 3000 ft (BPBM, OUMNH); Kona, 5000 ft (BMNH); Olaa (BPBM); Puna(BMNH); Waimea, 4000 ft (BPBM). Maui: Haleakala, 2000 ft (BPBM); West Maui Mts,Iao Valley (BMNH, BPBM). Oahu: Honolulu, 2000 ft (BMNH, BPBM); Honolulu Mts.,1800 ft (BPBM); Tantalus (BPBM, OUMNH).

Recent collections. Hawaii: Kipuka Nene, 2900 ft; Ocean View Estates, 4500 ft; PuuWaawaa, 4200 ft. Maui: Waihee Ridge Trail, 2000 ft; Puu Kukui Rd., 2300 ft; Kahoma,2100 ft. Lanai: Munro Trail, 3000 ft. Oahu: Aiea Trail, 1700 ft; Wiliwilinui Trail, 2000 ft;Papali Ridge, 1400 ft; Pahole Natural Area Reserve, 2000 ft; Palikea, 3000 ft; Puu Kaua,2600 ft. Kauai: Polihale, coast; Paaiki Valley, 2700 ft; Kahuamaa Flat, 3800 ft.

Other collections. Molokai: Pukoo, 1600 ft, 2♂ 1♀ 27-X-1913 (W. M. Giffard)(BPBM).

Flower records. Acacia, Bidens torta, Broussaisia arguta, Canavalia galeata,Chamaesyce multiformis, C. olowaluana, Claoxylon, Metrosideros, Myoporum (Kauai),Psychotria mariniana, Rubus rosifolius*, Scaevola chamissoniana, S. gaudichaudiana,S. procera, Sophora, Styphelia.

Remarks. Fullaway (1918:394, 396-397) pointed out the possible synonymy of H.connectens with H. koae and H. vicinus, now known as H. insulae. The three names areconsidered here to apply to one species. All the names were published in the same 1899publication by Perkins. The name H. connectens was chosen because it is based on the onlymale type and is not involved in homonymy. Each of the three names was originally describedfrom few specimens and only one sex: H. connectens based on a male from West Maui, H.vicinus based on a female from Hawaii and a junior homonym, and H. koae based on afemale from Oahu. We have seen bees of both sexes from Hawaii, Maui (both sexes fromthe type locality of H. connectens), Lanai (male only), Oahu (both sexes from the typelocality of H. koae), and Kauai. Allowing for geographic variation in color and punctation,both sexes from all the islands are otherwise similar. Males are similar in body size, widesupraclypeal area, punctation of the upper face, scape dilation and arch, propodeal sculpture,and terminalia. The female types of H. koae and H. vicinus and topotypical females of H.connectens are similar in body size and structural characters.

Based on small samples of mostly old specimens, bees from Oahu and Maui are moremelanic and with smoky wings in comparison to those from Kauai and Hawaii with slightlysmoky to clear wings. Males from Oahu and Maui have a wider supraclypeal area (W/Lratios 1.4-1.6, n = 11), only the clypeus marked with yellow (varying from a broad, fullmark to a small spot), and dark, unmarked legs (other than the usual pale fore tibia andrarely a basal yellow dot on the other tibiae). Males from Hawaii have a narrower


supraclypeal area (W/L ratios 1.2-1.3, n = 4) and more extensive yellow marking: clypeusand supraclypeal area yellow with or without an additional mark in the lower paraoculararea, and legs may have basal spots of yellow on the tibiae and basitarsi. Both males andfemales from Kauai may have marks on the pronotal lobes, and females may have marks onthe legs.

References. Hylaeus koae: Perkins, 1907a:43 (record: Oahu, Tantalus); 1910:603 (key);1911:727 (Blackburn had series of mixed species). Timberlake, 1920:362 (probableparasitoid: Nesencyrtus kaalae). Cockerell, 1926:309 (♂; record: Oahu, Mt. Tantalus).Beardsley, 1976:188, 209 (probable parasitoid: Coelopencyrtus kaalae). Nesoprosopisvicina: Perkins, 1907b:97 (record: Hawaii, Kilauea); 1910:603 (key).

Hylaeus (Nesoprosopis) crabronoides (Perkins)(Fig. 19)

Nesoprosopis crabronoides Perkins, 1899:94 (♀; Pl. 2, Fig. 2).Hylaeus (Nesoprosopis) crabronoides, Meade-Waldo, 1923:34 (list).Nesoprosopis crabronoides, Daly, 1994:116 (lectotype ♀ BMNH 17a 2752; Hawaii,

Kilauea; wing length, 3.7 mm).

DIAGNOSIS. Very small elongate bees with clear wings. Male with three separate largeyellow facial marks, yellow marks on the pronotal lobes and at the sides of the collar, andthe basal third of the hind tibiae yellow. Female with unique swollen frons and uniqueelongate mesosoma. See remarks below for comparison with sister species H. specularis.Males are distinguishable from H. specularis by the rather thick head and lack of animpunctate area above the antennal socket.

Distribution. Hawaii; montane wet forest.Male. Measurements: Malar space, 2; eye, 81:78:71:47; clypeus, 42×39; frons, 12×21:64;

interalveolar, 10:15:37:44, D 10; interocellar, 22:25:17:25, D 11; scape, 29×17; wing length,estimated 3.3 mm (Hawaii, Olaa Small Tract, 3800 ft, KM-00-132).

Head. Malar space very short (ratio about 0.2 DMO). Upper frons swollen. Upperparaocular area with pits less than 1 pit width apart, on frons smaller and crowded, distortedin subpolygonal shapes; a dark rhomboid spot in the center of the frons created by denseblack hairs and very dense punctation; small shiny area just below spot and above antennalsocket. Lower paraocular depression absent, surface flat. Frontoclypeal sulcus straight.Scape in frontal view slightly dilated, width more than half length, ratio about 0.6; lateraledge in frontal view nearly straight; slightly arched in lateral view with maximum arch nearmiddle of scape; underside of median edge with a groove; with long, curved, yellowishwhite hair along median edge. Vertex hair 8-14, pale brown, simple.

Mesosoma. Scutum with pits 1-3 pit widths apart, ground granular to minutely wrinkled,slightly lustrous; hair medium-length (mostly 6, scattered long hairs), pale brown.Mesepisternum with indistinct, shallow pits 1-2 pit widths apart, long white plumose hair;ground coriaceous, lustrous. Hypoepimeron weakly punctate, coriaceous, lustrous, nearly

HAWAIIAN HYLAEUS 69

flat. Basal area of propodeum almost entirely smoothly coriaceous, sometimes a few shortrugae at the front.

Metasoma. Elongate. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine,pale, appressed hairs spaced about one hair length apart; T7 hairs long, brown, erect.

Terminalia. Lateral process of S7 tapered, length greater than width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with elongate, narrow tips. Apicesof gonoforceps subequal to penis valves, inwardly pointed.

Color. Head and mesosoma black; metasoma blackish brown. Labrum and mandibleblack (one specimen with obscure yellowish red marks on mandible). Face with three separatemarks: clypeus largely yellow, with broad black basal and lateral margins; lower paraoculararea with triangular yellow spot along the eye to level of antennal socket; supraclypeal area

Fig. 19. Hylaeus crabronoides, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

A B C D

E F

1.0 mm


and scape unmarked. Pronotum with transverse collar stripe at sides; lobes mostly yellow.Legs dark brown; fore femur and tibia with large pale area; small yellow marks on fore andmid femur at very apex, fore tibia with diffuse stripe in basal half along pale area; mid tibiawith small spots at base and apex, sometimes connected by an obscure dorsal stripe; hindtibia yellow in basal third and sometimes narrowly at apex; fore and mid basitarsi yellowishbrown; hind tarsus dark brown.

Female. Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area andfrons strongly swollen, convex, clearly bulging out beyond eye at level of frontal sulcus inlateral view; upper paraocular area with minute round pits less than 1 pit width apart,ground shiny; frons with pits becoming crowded medially and partly distorted in shapenear the middle; frontal line distinct only near acme; small shiny area just above antennalsocket. Apex of clypeus shallowly emarginate. Dorsal end of facial foveae not extended tolevel of lateral ocelli. Vertex hair 6-10, pale brown, weakly plumose.

Mesosoma. Fore tarsus with hairs apically curved. Mesosoma elongate with dorsal surfacedepressed, long anterior area of propodeum nearly in same horizontal plane as the scutumand scutellum. Scutum with minute pits variable, distinct and round or poorly defined andobscured by sculpture of ground, 2 or more pit widths apart, ground slightly lustrous; hairextremely short (about 2, scattered longer ones up to 8), red-brown, simple. Scutellumwith ground smoothly coriaceous and lustrous. Pubescence of mesepisternum yellowishwhite. Metanotum coriaceous and slightly lustrous. Hypoepimeron impunctate, nearly flat.Basal area of propodeum with long anterior area, short, lineately rugose at front; browcoriaceous and lustrous.

Metasoma. Narrow, elongate. Disk of T2 impunctate, smoothly coriaceous, but polishedand shiny, with fine, pale, appressed hairs spaced about one hair length apart. T6 hairslong, dark brown, erect.

Color. Body dark brown. Face marks: clypeus unmarked or with an obscure subapicalmark (clypeus of type is unmarked); lower paraocular area with yellow triangular stripealong the eye to slightly above level of antennal socket. Pronotum with transverse collarstripe interrupted medially; lobes yellow. Legs dark yellowish brown except: fore tibia withlarge pale area edged at base by diffuse yellow stripe; femora with minute yellow marks atvery apices; fore and mid tibial bases with small basal yellow marks; hind tibia with basalyellow stripe up to nearly half the length of the segment. Wings clear.

Perkins’ collections. Hawaii: Kau, 4000 ft (BMNH); Kilauea (OUMNH); field notes#638, #533 (BMNH).

Recent collections. Hawaii: Kona Forest Unit of Hakalau National Wildlife Refuge,South Boundary Rd., 4700 and 5200 ft; Olaa Small Tract, 3800 ft.

Flower records. Cheirodendron, Myoporum.Remarks. Hylaeus crabronoides was described from a few females taken at Kilauea,

Hawaii, at 4000 ft. The very small, distinctive females (forewing length of type, 3.7 mm)have unique features: frons greatly swollen with the fovea not reaching the level of thelateral ocelli; and a narrow, elongate body with the mesosoma depressed such that thescutum, scutellum, metanotum, and anterior area of the propodeum are almost in the samehorizontal plane. It is closely related to H. specularis, in the company of which most of the

HAWAIIAN HYLAEUS 71

recent specimens have been collected. Females are easily distinguished by the swollenfrons, whereas males are told by the unusually thick head (about half as long as wide indorsal view, measured from the acme to the center of the occiput) and by the punctate areaabove the antennal sockets (versus smooth and shining). Both sexes can be separated fromthe much more common H. dumetorum; males by the arched scape, and females by themore lustrous upper frons and vertex.

References. Perkins, 1907b:97 (record: Hawaii, Kilauea); 1910:601 (key); 1913:lxxix(extreme morphology).

Hylaeus (Nesoprosopis) difficilis (Perkins)(Fig. 20)

Nesoprosopis difficilis Perkins, 1899:80-81 (♂♀; Pl. 2, Fig. 11).Hylaeus (Nesoprosopis) difficilis, Meade-Waldo, 1923:33 (list).Nesoprosopis difficilis, Daly, 1994:116 (lectotype ♂ BMNH 17a 159b; Hawaii, Kau,


DIAGNOSIS. Medium-sized bees with clear to smoky wings. Male face usually with ovalyellow mark or mark on clypeus only, hemisternites of S7 broadly triangular or with a shortprocess, process of S8 not dilated and apex simple or with very short bifurcations,gonoforceps subequal to penis valves. Female unmarked, hair of scutum long, of T6 medium-length and prostrate. Sister species of H. laetus; see remarks.

Distribution. Hawaii, Maui, Lanai, and Molokai; very widespread, found in lowlandand montane dry shrubland and dry to wet forest, and at the coast on Hawaii Island.

Male. Measurements: Malar space, 6; eye, 113:105:92:64; clypeus, 59×61; frons,20×23:82; interalveolar, 15:20:47:54, D 19; interocellar, 29:28:18111:26, D 15; scape,39×20; wing length, 4.0 mm (Hawaii, Kau, 4000 ft, BPBM 3311).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area with round,shallow pits 1 pit width apart or less (some specimens with pits indistinct or almost totallyobscured by minutely wrinkled to granular and dull ground; other specimens with relativelysmooth ground, lustrous); below punctured area and along the eye with dense, minute,parallel wrinkles, rather smooth and lustrous; frons with pits medially closer, becomingcontiguous, slightly distorted; fine frontal line sometimes present; shiny, median, depressedarea just above prominent acme. Lower paraocular depression distinct, but shallow.Frontoclypeal sulcus straight or nearly so. Scape in frontal view slightly dilated, widthabout half length, ratio about 0.5; lateral edge in frontal view nearly straight or slightlyconcave; strongly arched in lateral view with maximum arch slightly toward apex of scape,thickness of scape in lateral view more or less uniform in basal two-thirds, expanded inwidth in apical third; no groove on underside; with long, yellowish to brownish white,simple hairs. Vertex hair 14-24, brown, mixed simple and plumose.

Mesosoma. Scutum with shallow pits about 2 pit widths apart, pits sometimes indistinctdue to variable ground, minutely wrinkled to granular, dull to slightly lustrous; hair long(12-20), yellowish white to red-brown, plumose. Scutellum pits more distinct, ground


slightly lustrous to lustrous. Mesepisternum with punctation varying from barely discernibleto distinct, shallow, round pits, about 1 pit width apart, and long, yellowish white to white,plumose hair; ground minutely coriaceous, wrinkled or striate, slightly lustrous.Hypoepimeron impunctate, smooth to partly striate, slightly lustrous, nearly flat. Metanotumdull. Basal area of propodeum of some specimens lineately rugose to brow, 2-4 ruga widthsapart varying to irregular, reticulately rugose to brow; often shiny between rugae; browgranularly coriaceous, dull to slightly lustrous; other specimens with lineate rugae notreaching brow, brow smoothly coriaceous and lustrous.

Metasoma. Disk of T2 impunctate; smoothly coriaceous, shiny, with bare areas or pale,short, appressed hairs spaced more than a hair length apart. T7 hairs short, golden to darkbrown, appressed.

Fig. 20. Hylaeus difficilis, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.

0.5 mm

A B C D

E F

1.0 mm

HAWAIIAN HYLAEUS 73

Terminalia. Lateral process of S7 tapered, acute, with length equal to width ofhemisternite, and brown sclerotized color throughout; overall shape variable, broadlytriangular or slightly L-shaped. Median process of S8 linear and with apex simple and hairyor a slight, triangular dilation at tip with short, slender, cuticular bifurcations with hairs(the hairs at tip of process may look like long bifurcations). Apices of gonoforceps subequalto penis valves.

Color. Labrum and mandible blackish brown to black. Face with a single large spot:clypeus entirely yellow or nearly so; lower paraocular area yellow along sulcus belowantennal socket but not contacting eye, in some specimens reduced to irregular stripealong suture; supraclypeal area to acme yellow (the supraclypeal and paraocular areasunmarked in some specimens); scape unmarked. Pronotum unmarked. Legs dark brown toblack and unmarked except pale area on fore tibia, or with some or all of the followingyellow marks: small spot at apex of fore femur, basal spot or stripe on fore tibia, basalsmall spot on mid tibia, basal stripe up to half length of hind tibia, all basitarsi with a stripe.Wings clear to smoky. Specimens from coastal populations and rare montane individualsmay have marks on the pronotal collar and lobes, and on the labrum; such specimensusually have the maximum leg markings and broad paraocular marks similar to those of H.anthracinus.

Female. Head. Malar space medium (ratio about 0.5 DMO). Upper paraocular areawith indistinct, round, shallow pits 1 pit width apart or less; ground minutely wrinkled togranular, dull; frons with pits medially closer, becoming contiguous, slightly distorted; finefrontal line sometimes present. Apex of clypeus shallowly emarginate. Dorsal end of facialfovea extended to level of lateral ocellus. Vertex hair 12-20, brown, plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with shallow pits about 2 pitwidths apart, pits sometimes indistinct due to variable ground, minutely wrinkled to granular,dull to slightly lustrous; hair long (10-14), yellowish white to red-brown, plumose. Scutellumpits more distinct, ground slightly lustrous. Mesepisternum with punctation varying frombarely discernible to distinct, shallow, round pits, about 1 pit width apart, and long, yellowishwhite to white, plumose hair; ground minutely coriaceous, wrinkled or striate, slightlylustrous. Hypoepimeron impunctate, smooth to partly striate, slightly lustrous, nearly flat.Metanotum dull. Basal area of propodeum lineately rugose to brow, 2-4 ruga widths apartvarying to irregular, reticulately rugose to brow; brow granularly coriaceous, dull to slightlylustrous.

Metasoma. Disk of T2 impunctate; smoothly coriaceous, shiny, with bare areas or pale,short, appressed hairs spaced more than a hair length apart. T6 hair medium-length, goldento dark brown, prostrate.

Color. Head and mesosoma black; metasoma dark brown. Legs dark brown to black;unmarked. Face and pronotum usually unmarked; coastal specimens sometimes with asmall, irregular, subapical mark on clypeus or a small basal mark on mid tibia.

Perkins’ collections. Hawaii: Kau, 4000 ft (BMNH, BPBM); Kilauea, 4000 ft (BPBM,OUMNH); Kona, 4000 ft (BMNH, BPBM), plateau, 4000 ft (BPBM), 5000 ft (BMNH,BPBM). Maui: Haleakala crater (BPBM); West Maui Mts. Iao Valley (BMNH, BPBM).Lanai: Halepaakai, 3000 ft (BMNH); “Lanai,” 2000 ft (BMNH, BPBM); Mts. Koele,


2000 ft (BMNH). Molokai: Molokai Mts. (BPBM), lower slopes (BPBM), 3000 ft (BMNH,BPBM), 4000 ft (BPBM).

Recent collections. Hawaii: Devastation Trail, 3700 ft; Kipuka Ki, 4200 ft; KipukaNene, 2900 ft; 0.9 mi. SW of Mauna Loa Rd. (near Kipuka Maunaiu), 6000 ft; KipukaAlala, 5700 ft; Puu Huluhulu (N. Hilo), 6700 ft; Puu Waawaa, 4100, 5500, and 6700 ft;Ocean View Estates, 2400 ft; Kaalualu, coast; South Point, coast; Kaloko-HonokohauNational Historic Park, coast. Maui: Kahoma, 2100 ft; Puu Kukui Rd., 2300 ft; Hanaula,above Puu Anu, 915 m; Waikamoi Preserve boardwalk, 6400 ft; Kaupo Trail, 6000 ft;Koolau Gap, 6200 ft; Puu Keokea, 7100 ft; above Hosmer Grove, 2100 m; Kalepeamoa,2835 m; Kilohana Pali, 2800 m; Kipahulu Valley, Mauka Ridge, 1980 m; Oili Puu, 2050 m.Lanai: Munro Trail, 2800 and 3000 ft. Molokai: Kamakou Rd., 3200 ft; Puu Kolekole,3400 and 3900 ft.

Flower records. Acacia, Argyroxiphium sandwicense, Bidens menziesii, Chamaesycecelastroides, C. olowaluana, Cheirodendron, Dodonaea, Dubautia ciliolata, D. menziesii,D. scabra, Metrosideros, Myoporum, Santalum paniculatum, Sapindus saponaria, Scaevolasericea, Schinus terebinthifolius*, Sophora, Styphelia, Tournefortia*, Vacciniumreticulatum.

Remarks. Hylaeus difficilis (Maui Nui and Hawaii) and H. laetus (Kauai to Hawaii) arebroadly sympatric sister species that are closely similar in sculpture. Each species has variableyellow markings such that some specimens in each may be similar in appearance. In hiskey, Perkins (1910:602) distinguished the males on the basis of the apex of the medianprocess of S8 (longish bifurcation in H. laetus, not so in H. difficilis; but sometimes H.difficilis may have a moderately long bifurcation) and the pubescence on the mesonotum(rather long in H. difficilis; short in H. laetus). Both species have a similarly marked face,including the clypeus, lower paraocular areas, and supraclypeal area in one large yellowmark or only the clypeus. Hylaeus laetus may or may not have an apical yellow spot on theprothoracic lobes and usually has basal stripes on the tibiae, up to half the length of thehind tibia, and yellow basitarsi. Hylaeus difficilis almost never has a spot on the lobes andusually has less yellow markings on the legs or none, but some specimens of H. difficilismay have similar leg marks. Coastal specimens from Hawaii may have marks on theprothoracic collar and lobes, as well as extensive marks on the legs. Hylaeus difficilis hasthe lateral process of S7 shorter, the apex narrow, and of a sclerotized brown color. In H.laetus, the lateral process is broad, translucent apically, and longer than the width of thehemisternite. Females can be separated by the shorter scutum hair of H. laetus.

In more montane areas H. difficilis may occur with H. volcanicus and H. nivicola, bothof which have longer malar spaces. Males can also usually be told from the former speciesby the simple process of S8; while it may be prolonged into bifurcations that can bemoderately long, those of H. volcanicus are always much longer. Females can be moredifficult to separate, as the malar space of H. volcanicus is variable. The smaller bee H.coniceps also can be found with them. The males are distinctive, but the females are similarto H. difficilis and may be distinguished from that species by the smaller malar area and thelonger, erect, black hairs of T6.

HAWAIIAN HYLAEUS 75

References. Perkins, 1899:105 (host of cleptoparasites: N. sphecodoides, N. volatilis);1907a:43 (record: Oahu, Tantalus); 1907b:97 (record: Hawaii, Kilauea); 1910:602 (key);1911:727 (Blackburn had series of mixed species); 1913:79, 82 (biology, biogeography).Fullaway and Giffard, 1919:51 (record: Maui, Olinda, 4200 ft). Swezey and Williams,1932:181 (record: Hawaii, Nauhi Gulch; flower: Sophora, Dubautia [referred to asRaillardia]). Swezey, 1954:205 (flower: Sophora chrysophylla; record: Hawaii, Nauhi,8000 ft). Suehiro, 1986:60 (record: Hawaii, Kona).

Hylaeus (Nesoprosopis) dimidiatus (Perkins)(Fig. 21)

Nesoprosopis dimidiata Perkins, 1899:96 (♂♀).Hylaeus (Nesoprosopis) dimidiatus, Meade-Waldo, 1923:34 (list).Nesoprosopis dimidiata, Daly, 1994:116-117 (lectotype ♂ BMNH 17a 149; Hawaii,

Kona, 4000 ft).

DIAGNOSIS. Small bees with clear wings, propodeum lineately rugose in basal third,nearly smooth to brow. Male with lower face yellow up to level of antennae or very narrowlyabove, pronotal collar at sides and lobes yellow, tibiae and tarsi marked with yellow. Lackingthe scape mark of H. hula, and the mandibular mark of H. kona. Similar to specimens of H.pele with large face marks, but lacking a groove on the underside of the scape and with thepropodeum sloping. Female with yellow marks on the lower paraocular areas, pronotallobes and sides of collar, base of front and mid tibiae, and rather widely banded on hindtibiae; differing from similar species by the evenly spaced pits of the frons and slopingpropodeum.

Distribution. Hawaii; montane mesic and dry forest.Male. Measurements: Malar space, 3; eye, 99:95:86:59; clypeus, 49×52; frons, 18×25:72;

interalveolar, 18:17:40:48, D 18; interocellar, 28:25:18:25, D 14; scape, 34×22; wing length,3.3 mm (H. dimidiatus lectotype).

Head. Malar space short (ratio about 0.3-0.4 DMO). Upper paraocular area and fronsto middle with very indistinct shallow pits about 1 pit width apart, pits obscured by minutelywrinkled ground, dull; frontal line indistinct; large impunctate, minutely wrinkled area aboveantennal socket. Lower paraocular depression absent. Frontoclypeal sulcus straight to slightlyarched. Scape in frontal view slightly dilated, width more than half length, ratio about 0.5-0.6, triangular, greatest width near apex; lateral edge in frontal view nearly straight;moderately arched in lateral view with maximum arch near middle of scape; no groove onunderside; with medium-length, white, plumose hair. Vertex hair 10-16, yellowish white,plumose.

Mesosoma. Scutum with very indistinct shallow pits about 2 pit widths apart, pitsobscured by minutely wrinkled ground, slightly lustrous; hair short (4-8), pale, plumose.Scutellum with larger, more distinct pits, lustrous. Mesepisternum with very indistinct pitsabout 2 pit widths apart, and long, yellowish white, plumose hair; ground smooth, lustrous.


Hypoepimeron impunctate, slightly swollen, shiny. Metanotum minutely wrinkled, dull toslightly lustrous. Basal area of propodeum rather short and sloping with lineate rugae inbasal third, spaced 2-3 rugae apart, shiny between rugae; smoothly coriaceous, lustrous toand including brow.

Metasoma. Disk of T2 impunctate, smooth, shiny, with fine, pale hairs spaced morethan a hair length apart. T7 hair short to medium, red-brown, erect.

Terminalia. Lateral process of S7 tapered, pale, length greater than width of hemisternite.Median process of S8 dilated, apex bifurcate with short, rounded tips. Apices of gonoforcepssubequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face with one large mark: clypeusentirely yellow or with very narrow lateral margins black; lower paraocular area yellow to

Fig. 21. Hylaeus dimidiatus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

A B C D

F

1.0 mm

E

HAWAIIAN HYLAEUS 77

level of antennal socket or very narrowly above; supraclypeal area yellow to below antennalsockets; scape unmarked. Pronotum with transverse collar stripe interrupted medially; lobesyellow at least in part. Fore leg dark brown except: fore femur with apex yellow; fore tibiawith dorsal yellow stripe up to full length; fore basitarsus entirely yellow and at least thebasal mediotarsus yellow. Mid and hind legs dark brown to black except: mid femur withapex yellow; mid and hind tibiae with basal and apical yellow marks; basitarsi entirelyyellow; at least the basal mediotarsus yellow. Wings clear.

Female. Head. Malar space short (ratio about 0.5 DMO). Upper paraocular area andfrons with shallow, evenly spaced pits, about 1 pit width apart even near middle; groundgranular and dull. Frontal line obscure. Apex of clypeus slightly emarginate. Dorsal end offacial fovea reaching level of lateral ocellus. Vertex hairs 10-14, yellowish white, plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with shallow pits 2-3 pitwidths apart, ground minutely granular and slightly lustrous; hair very short (2-4), palebrown, plumose. Scutellum similar, slightly more lustrous. Metanotum roughened, dull.Mesepisternum with shallow pits 1-2 pit widths apart, ground minutely granular, lustrous;with medium-length, pale brown, weakly plumose hair. Hypoepimeron impunctate, lustrous,hardly swollen. Basal area of propodeum with a few very short rugae at base, the remainderincluding the brow smoothly coriaceous, lustrous; brow rounded, not prominent.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,prostrate hairs spaced two hair lengths apart or more. T6 hairs long, brown, erect.

Color. Head and mesosoma black; metasoma dark brown. Face marked with yellow:paraocular area with yellow spots, sometimes reduced or absent. Pronotum with marks onlobes and interrupted transverse collar stripe. Legs black except: front tibia with large palearea and a dorsal yellow stripe one-third to half the length; mid tibia with basal quarteryellow; hind tibia with basal half yellow. Wings slightly smoky.

Perkins’ collections. Hawaii: Kona, 4000 ft (BMNH).Recent collections. Hawaii: Puu Waawaa, 4100 ft; Kipuka Alala, 5800 ft.Flower records. Chamaesyce olowaluana.Remarks. This is a rare species apparently restricted to dry to mesic forest in Kona; it

and the sympatric H. akoko and H. kona are the only montane species from Hawaii thathave never been collected at Kilauea. Despite appearances, genetic data clearly shows it tobe the sister species of H. connectens. The identity of the female has been the subject ofsome confusion. Fullaway (1918) described it from a specimen caught with the male atKahuku, believing Perkins’ (1899) original association to be incorrect. Perkins stated (1899)that while he believed his specimen had come from the same place as the male, the localitylabel had been lost. The latter description appears to match H. pele, which occurs with H.dimidiatus in Kona; it is the only species with a smooth propodeum that has the apicalhairs of the metasoma somewhat pale. The specimens on which the two descriptions werebased could not be found. Recent specimens have been genetically confirmed, and agreewith Fullaway’s description.

References. Fullaway, 1918:395-396 (♀; record: Hawaii, Kahuku, Kau). Suehiro,1986:60 (record: Hawaii, Kona).


Hylaeus (Nesoprosopis) dumetorum (Perkins)(Fig. 22)

Nesoprosopis dumetorum Perkins, 1899:92-93 (♂♀).Hylaeus (Nesoprosopis) dumetorum, Meade-Waldo, 1923:34 (list).Nesoprosopis dumetorum, Daly, 1994:117 (lectotype ♂ BMNH 17a 163a; Hawaii, Puna,


DIAGNOSIS. Small, elongate bees with clear wings. Males with three separate yellowmarks on face, variably marked on pronotum and legs; distinguishable by the unarchedscape. Females marked with narrow yellow stripes along the eyes, marks on the pronotalcollar and lobes, and the basal quarter or less of the hind tibiae is yellow. Sister species ofH. angustulus.

Distribution. Hawaii Island; widespread and common in montane wet and mesic forest.Male. Measurements: Malar space, 3; eye, 101:95:87:64; clypeus, 51×54; frons,

18×24:73; interalveolar, 18:20:44:49, D 17; interocellar, 20:28:13:26, D 13; scape, 37×20;wing length, 4.0 mm (Hawaii, Kau, 4000 ft, BPBM 3342).

Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular area and fronswith pits less than 1 pit width apart, crowded and distorted in subpolygonal shapes; a darkrhomboid spot at the middle due to dense black hairs and very dense punctation (may beless distinct in soiled specimens); shiny area just below spot and above antennal socket.Lower paraocular depression absent, surface flat. Frontoclypeal sulcus straight or nearlyso. Scape in frontal view slightly dilated, width about half length, ratio about 0.5; lateraledge in frontal view nearly straight; median edge gently rounded, pubescent; underside ofmedian edge without a narrow groove, but with a broad depression with granular sculpture;almost flat in lateral view. Vertex hair 10-18, brown, simple.

Mesosoma. Scutum with indistinct, minute, shallow pits more than 2 pit widths apart,ground dull coriaceous; hair short (4-6 with scattered longer hairs), red-brown, simple.Scutellum with slightly larger pits 1-2 pit widths apart, ground slightly lustrous.Mesepisternum with minute, indistinct pits 2-4 pit widths apart, and long, yellowish white,plumose hair; ground smoothly coriaceous, slightly lustrous to lustrous. Hypoepimeronimpunctate, slightly swollen. Metanotum slightly lustrous. Basal area of propodeum withvariable sculpture, sparse, short, linear or reticulate rugae at the front or nearly glabrous;brow smoothly coriaceous, lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart. T7 hair long, black, erect.

Terminalia. Lateral process of S7 parallel-sided, pale, length greater than width ofhemisternite. Median process of S8 strongly dilated; apex bifurcate with broad tips. Apicesof gonoforceps subequal to penis valves.

Color. Head and mesosoma reddish brown to black; metasoma dark reddish brown todark blackish brown. Labrum and mandible blackish brown to black. Face with three separatemarks: clypeus with a subapical, irregular yellow spot (rarely absent); lower paraoculararea with triangular yellow mark along the eye below antennal socket; supraclypeal areaand scape unmarked. Pronotum usually unmarked, sometimes with a mark on the pronotallobe and rarely the collar. Legs reddish brown to black except: fore tibia with pale area,with or without dorsal yellow stripe and fore femur with yellow mark at very apex; mid and

HAWAIIAN HYLAEUS 79

hind tibiae variable, with or without an obscure yellowish mark at extreme base. Wingsclear to smoky.

Female. Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area andfrons convex; upper paraocular area with nearly round pits less than 1 pit width apart,ground lustrous to shiny; frons with pits becoming crowded below the median ocellus andpartly distorted in shape, and near the middle a few, fine, longitudinal carinulae; frontal linedistinct only near acme; small shiny area just above antennal socket. Apex of clypeusshallowly to distinctly emarginate. Dorsal end of facial foveae extended just to level oflateral ocelli. Vertex hair 8-12, brown, simple.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with indistinct, minute, shallowpits more than 2 pit widths apart, ground dull to very dull; hair short (4-6) brown, weakly

Fig. 22. Hylaeus dumetorum, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

0.5 mm

A B C D

E F

1.0 mm


plumose. Scutellum with pits 1-2 pit widths apart, ground slightly lustrous in some specimens.Mesepisternum with minute, indistinct pits 2-4 pit widths apart, and long, yellowish white,plumose hair; ground smoothly coriaceous, slightly lustrous to lustrous. Hypoepimeronimpunctate, slightly swollen. Metanotum dull. Basal area of propodeum with sparse, obscure,lineate rugae at very front; brow smoothly coriaceous, lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart. T6 hair long, dark brown, erect.

Color. Head and mesosoma black; metasoma dark brown. Face marks: lower paraoculararea with yellow stripe of variable width along the eye below antennal socket; clypeus andsupraclypeal area unmarked. Pronotum with transverse collar stripe interrupted medially;lobes yellow at least in part. Tegula with or without yellow spot. Legs reddish brown todark brown except: fore tibia with pale area combined with short yellow stripe at base; midand hind tibia with yellow spot at base. Wings clear to smoky.

Perkins’ collections. Hawaii: Hilo, 1800 ft (BMNH, BPBM); Kau, 4000 ft (BMNH,BPBM); Kilauea (BPBM, OUMNH); Olaa (BPBM); Puna, 2000 ft (BMNH, BPBM).

Recent collections. Hawaii: Byron’s Ledge, 3800 ft; Halemaumau Trail, 3800 ft; KonaForest Unit of Hakalau National Wildlife Refuge, South Boundary Rd., 4500-5200 ft;Kipuka Ki, 4200 ft; Kipuka Puaulu, 4000 ft; Kau Forest Reserve, Mountain House Rd.,3000 ft; Upper Waiakea FR, 3700 ft; Upper Hamakua Ditch, 3700 ft.

Flower records. Alphitonia ponderosa, Buddleia asiatica, Cheirodendron, Crocosmia× crocosmiiflora*, Dubautia scabra, Fragaria vesca*, Hedyotis terminalis, Ilex anomala,Metrosideros, Myoporum, Myrica faya*, Pisonia brunoniana, Psychotria hawaiiensis,Rubus rosifolius*, Styphelia.

Remarks. The most common wet and mesic forest species on Hawaii. Distinguishablefrom the other common species, H. coniceps, by the flat scape, short hairs of the scutum,and reduced rugosity of the propodeum. In males, the facial marks are distributed differently:H. coniceps often has the paraocular marks shorter than the clypeal mark, and if the marksare reduced the former disappear first; H. dumetorum has the paraocular marks longerthan the clypeal (one melanic specimen lacks the clypeal mark). Females resemble H.specularis, but have the upper frons only slightly lustrous, with a narrower dull area at themiddle, and the impunctate area laterad of the lateral ocellus distinctly coriaceous ratherthan almost smooth and brightly reflective. Sister species of H. angustulus of Maui Nui.

References. Perkins, 1907b:97 (record: Hawaii, Kilauea); 1910:602 (key); 1913:82(biogeography). Fullaway, 1918:395 (record: Hawaii, Kilauea).

Hylaeus (Nesoprosopis) facilis (F. Smith)(Fig. 23)

Prosopis facilis F. Smith 1879a:683; 1879b:21 (♂♀).Nesoprosopis facilis, Perkins, 1899:77-78 (♂♀; Pl. 2, Figs. 1, 8).Hylaeus (Nesoprosopis) facilis, Meade-Waldo, 1923:33 (list).Prosopis facilis, Daly, 1994:119-120 (lectotype ♂ BMNH 17a-154ab; Smith gave Maui

as type locality, but Blackburn, 1886:143 and 1887:203, stated true locality is Oahu,Pauoa Valley).

HAWAIIAN HYLAEUS 81

DIAGNOSIS. Medium-sized bees with smoky wings. Male face with oval yellow mark,legs not marked with yellow or obscurely so, unusual lateral process of S7 long and broad,median process of S8 not dilated and with long narrow tips, unusual gonoforceps expandedapically beyond the penis valves and with pubescence not extended around apex to innerangle. Closely related to H. chlorostictus and H. simplex.

Distribution. Maui, Molokai, Lanai, Oahu; coast to mountains, dry shrubland to wetforest. Only one recent collection, from Maui.

Male. Measurements: Malar space, 5; eye, 142:114:108:72; clypeus, 69×71; frons,28×22:92; interalveolar, 12:25:49:62, D 22; interocellar, 29 29:22:31, D 17; scape, 55×28;wing length, 4.5 mm (H. facilis lectotype).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area and frons ratherflat, with long plumose hair, and with indistinct, mostly round, shallow pits 1 pit width orless apart even at the middle, ground minutely wrinkled, the wrinkles more or less paralleltransversely across the middle and longitudinally at sides along eye, slightly lustrous; median,impunctate, lustrous area above antennal sockets, slightly depressed and sometimes withone or two minute holes at middle. Lower paraocular depression distinct, but shallow.Frontoclypeal sulcus slightly arched. Scape in frontal view slightly dilated, width half length,ratio about 0.5; lateral edge in frontal view nearly straight or slightly concave; stronglyarched in lateral view with maximum arch near middle of scape; no groove on underside;with short to medium-length, pale brown, weakly plumose hair. Vertex hair 20-30, brown,plumose.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground slightly granular and slightlylustrous; hair medium-length (12-20), brownish white, plumose. Scutellum with slightlylarger pits, smoothly coriaceous, lustrous. Mesepisternum with distinct to indistinct, shallow,round pits, about 1 pit width apart, and long, yellowish white, plumose hair; ground minutelycoriaceous or striate, slightly lustrous. Hypoepimeron impunctate, smooth, lustrous, slightlyswollen. Metanotum dull. Basal area of propodeum lineately rugose to brow, shiny betweenrugae; at brow and below granular or somewhat smoothly coriaceous, dull to slightlylustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with short, fine, pale,appressed hairs spaced about one hair length apart or more. T7 hair medium-length, brown,prostrate.

Terminalia. Lateral process of S7 broad, sword-shaped, length much greater than widthof hemisternite. Median process of S8 linear, apex bifurcate with long, filamentous, denselyhairy tips. Apices of gonoforceps exceeding penis valves, conspicuous even when retracted.

Color. Labrum and mandible blackish brown to black. Face with a single large spot notextended at the sides above the antennal sockets and with fine black sulci in “H” shape:clypeus entirely yellow or nearly so; lower paraocular area with a rather narrow yellowstripe below antennal socket and along sulci; supraclypeal area yellow to acme or below;scape unmarked. Pronotum unmarked. Legs dark brown to black except: fore tibia, andsometimes fore femur, with pale areas that may be yellowish along dorsal margins. Wingslight smoky.

Female. Head. Malar space medium-length (ratio about 0.6 DMO). Upper paraoculararea with shallow, nearly round pits about 1 pit width apart, ground smooth, lustrous;frons with pits medially smaller, contiguous, slightly distorted; ground minutely wrinkled,


slightly lustrous; faint frontal line in some bees. Apex of clypeus shallowly emarginate.Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 20-30, brown,plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with pits 1-2 pit widths apart,ground slightly granular and slightly lustrous; hair medium-length (12-24), pale yellowishbrown, weakly plumose. Scutellum with slightly larger pits, smoothly coriaceous, lustrous.Mesepisternum with indistinct, shallow, round pits, about 1 pit width apart, and long,yellowish white, plumose hair; ground minutely coriaceous, slightly lustrous. Hypoepimeronimpunctate, smooth, slightly lustrous, nearly flat. Metanotum dull. Basal area of propodeumlineately rugose to brow, shiny between rugae; below brow smoothly coriaceous, slightlylustrous.

Fig. 23. Hylaeus facilis, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.

0.5mm

A B C D

E F

1.0 mm

HAWAIIAN HYLAEUS 83

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with short, fine, pale,appressed hairs spaced about one hair length apart. T6 hair medium, brown, prostrate.

Color. Head and mesosoma black; legs and metasoma dark brown; unmarked. Wingsdark smoky.

Perkins’ collections. Maui: Haleakala, 5000 ft (BPBM); Iao Valley (BMNH, BPBM);“Maui” (BMNH, BPBM); W. Maui Mtns, 2000 ft (BMNH, BPBM, OUMNH); Wailuku,on sandy isthmus (OUMNH). Lanai: center of island, forest above Koele, 2000 ft(OUMNH); Halepaakai, 3000 ft (BPBM); “Lanai,” 2000 ft (BMNH, BPBM, OUMNH);Koele Mts. >2000 ft (BPBM). Molokai: E. mountains, 2000 ft (OUMNH); Molokai Mts.,3000 ft (BMNH, BPBM), >3000 ft (BPBM), 3500 ft (BPBM). Oahu: Honolulu (BMNH,BPBM), 2000 ft (BMNH, BPBM); Honolulu Mts. 1500 ft (OUMNH); Makana, Waianae,3000 ft (BPBM); Kaala Mts., 2000 ft (BMNH, BPBM); “Oahu” (BPBM); Tantalus (BPBM);Waianae Mts., 2000 ft (BMNH, BPBM).

Recent collections. Oahu: Poamoho Trail, 6-XII-1975, G. Nishida. Maui: Kokomo,nr. Makawao 1500 ft.

Flower records. None.Remarks. Colors of the female integument and pubescence are based on old specimens.

Apparently the most common dryland species in Perkins’ time, H. facilis and its sisterspecies H. simplex seem to have almost completely disappeared, despite continuedflourishing of H. difficilis and H. laetus in these habitats. Although it may (like other long-uncollected species) survive in numbers at unvisited locales, there is no question that it hasbeen extirpated from much of its range. Females of this species and H. simplex tend to beslightly larger and with slightly shorter hair than H. difficilis, but cannot be reliablydistinguished. However, given the steep decline in the former two species, any femalekeying to these three can be assumed to be H. difficilis unless an associated male is found.

References. Blackburn and Kirby, 1880:85 (records). Blackburn and Cameron, 1886:142-143, 148 (♂♀; key); 1887:202-203, 208 (♂♀; key). Dalla Torre, 1896:22 (list). Perkins,1899:105 (host of cleptoparasite: N volatilis); 1907a:43 (record: Oahu, Tantalus); 1910:601(key); 1911:727 (Blackburn had series of mixed species); 1913:64, 79 (biology,biogeography). Fullaway and Giffard, 1919:52 (record: Maui, Lahaina). Swezey and Bryan,1929:297, 310-312 (nest: in dead twig, in dead twig of Broussaisia arguta, in hollow twigof Clermontia grandiflora, in beetle burrow in dead stem of Smilax sandwicensis; record:Molokai, E. Kawela, Kamiloloa, Waikolu). Krauss, 1944:94 (record: Molokai, WaikoluValley; Maui, Hana). Swezey, 1954:203 (nest: in dead stem of Smilax; record: Molokai,Kainalu).

Hylaeus (Nesoprosopis) filicum (Perkins)(Fig. 24)

Nesoprosopis filicum Perkins, 1911:722 (♂; Hawaii, Kilauea).Hylaeus (Nesoprosopis) filicum, Meade-Waldo, 1923:35 (list).Nesoprosopis filicum, Daly, 1994:116 (type not located).


DIAGNOSIS. Medium-sized bees with light smoky wings. Both sexes with unusualextremely dense punctation across the entire upper paraocular area and frons. Male facewith three separate marks, those of paraocular areas extended above antennal sockets innarrow vittae; scape moderately dilated and strongly arched, with an apical knoblikeexpansion in lateral view; pronotum unmarked; process of S8 strongly dilated and withbroad, round tips. Female with small paraocular marks and yellow marks on pronotal collarbut not lobes. See diagnosis and remarks under H. mimicus.

Distribution. Hawaii; montane wet to mesic forest.Male. Measurements: Malar space, 5; eye, 105:102:97:59; clypeus, 59×54; frons,

17×24:73; interalveolar, 17:23:44:49, D 18; interocellar, 28:28:18:28, D 15; scape, 39×27;wing length, 4.2 mm (Hawaii, Kilauea, 4000 ft, BPBM 3385).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area and frons withpits less than 1 pit width apart, crowded and distorted in subpolygonal shapes; a darkrhomboid spot at the middle created by dense black or dark brown hair and very densepunctation; shiny area just below spot and above antennal socket. Lower paraoculardepression absent. Frontoclypeal sulcus straight to slightly arched. Impunctate area lateradof lateral ocellus shining. Scape in frontal view moderately dilated, width more than halflength, ratio about 0.7; lateral edge in frontal view nearly straight; strongly arched in lateralview (under surface much more strongly arched than the upper) with maximum arch abouttwo-thirds along, with an abrupt thickening at the apex; underside of median edge with agroove; with long, simple, brown hair. Vertex hair 16-24, brown, simple.

Mesosoma. Scutum with indistinct, shallow pits about 1 pit width apart, ground granular,dull; hair long (12-20), simple, brown. Scutellum with slightly larger pits, dull.Mesepisternum with shallow pits, about 1 pit width apart, and long, yellowish white, plumosehair; ground minutely coriaceous, slightly lustrous. Hypoepimeron impunctate, smooth,lustrous, swollen. Metanotum dull. Basal area of propodeum with a rather long, slightlyconcave anterior area; sparse, weak, lineate rugae at base; brow smoothly coriaceous andlustrous, prominent.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, dark, erecthairs spaced about one hair length apart. T7 hair long, dark brown, erect.

Terminalia. Lateral process of S7 parallel-sided, length subequal to width of hemisternite.Median process of S8 strongly dilated, apex bifurcate with broad, round tips. Apices ofgonoforceps subequal to penis valves.

Color. Head and mesosoma dark brown to black; metasoma dark brown. Labrum andmandible blackish brown to black. Face with three separate marks: clypeus with variableapical mark, extending up to half length of plate; paraocular area yellow, extended at sidesalong the eye above antennal socket as narrow vitta; supraclypeal area unmarked; scapeunmarked. Pronotum unmarked. Legs dark brown to black except fore tibia with pale area,apex of fore femur and all tibiae with or without a trace of yellow at base. Mid and hindtibia variable, with or without an obscure trace of yellow at base. Wings clear to lightsmoky.

Female. Head. Malar space long (ratio about 0.7 DMO). Upper paraocular area withmostly round pits less than 1 pit width apart; frons with pits becoming crowded medially,

HAWAIIAN HYLAEUS 85

distorted, and obscured along middle by longitudinal wrinkles; slightly lustrous ground.Apex of clypeus distinctly emarginate. Dorsal end of facial foveae not extended to level oflateral ocelli. Vertex hair 12-24, brown, simple.

Mesosoma. Fore tarsus with hair apically curved. Scutum with shallow, indistinct pits 1-2 pit widths apart, ground granular dull to slightly lustrous; hair medium-length (8-16)simple, pale brown. Scutellum with pits slightly farther apart, slightly lustrous.Mesepisternum with barely discernible pits, about 2-3 pit widths apart, and long, yellowishwhite, weakly plumose hair; ground smooth, lustrous. Hypoepimeron impunctate, smooth,lustrous, nearly flat. Metanotum slightly lustrous. Basal area of propodeum with sparse,weak, short rugae at very front, otherwise glabrous to brow; brow smoothly coriaceousand lustrous. Hypoepimeron impunctate, nearly flat.

0.5 mm

A B C D

E F

1.0 mm

Fig. 24. Hylaeus filicum, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.


Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, almost polished; fine,pale, semierect hair about one hair length apart. T6 hair long, brown, erect.

Color. Head and mesosoma dark brown to black; metasoma dark brown. Face marks:clypeus and supraclypeal area usually unmarked (clypeus rarely with an obscure subapicalspot); lower paraocular area with narrow, irregular yellow stripe along the eye up to levelof antennal sockets. Pronotum with transverse collar stripe interrupted medially; lobesunmarked. Legs dark brown except fore tibia with or without pale area and obscure basalyellow spot. Wings slightly smoky.

Perkins’ collections. None. Type collected later by Perkins at Kilauea, Hawaii.Recent collections. Hawaii: Puu Waawaa, 4100 ft; Kona Forest Unit of Hakalau National

Wildlife Refuge, South Boundary Rd., 5200 ft.Flower records. Chamaesyce olowaluana, Myoporum.Remarks. Types have not been located. Despite the close resemblance of the males to

H. mimicus, the females have much less facial coloring, and the species is probably moreclosely related to H. haleakalae of Maui Nui. Females in older Kilauea collections wereoften placed with males of H. hula, although the two species were apparently not collectedtogether in that area.

References. Fullaway, 1918:396 (variation).

Hylaeus (Nesoprosopis) finitimus (Perkins)Nesoprosopis finitima Perkins, 1899:100 (♂♀).Hylaeus (Nesoprosopis) finitimus, Meade-Waldo, 1923:34 (list).Nesoprosopis finitima, Daly, 1994:116 (lectotype ♀ BMNH 17a 168; Kauai, coast at

Makaweli; wing length, 5.0 mm).

DIAGNOSIS. Medium-sized bees with clear wings (only female type available for study).Perkins (1899:100) described the male H. finitimus only as having “the face shaped andcolored as in H. longiceps but with the punctation of the scutum and scutellum distinctlyfiner.” In his key (1910:602) H. finitimus has the face shorter and the supraclypeal areaabout as long as wide compared to H. longiceps, which has the face longer and the apicalmargin of the supraclypeal area much shorter than the sides. Female black and unmarked.

Distribution. Kauai; coast. No recent collections.Male. No type or other specimens available (see diagnosis).Female. Head. Malar space short (ratio about 0.4 DMO). Apex of clypeus shallowly

emarginate. Dorsal end of facial fovea extended to level of lateral ocellus. Upper paraoculararea with round pits about 1 pit width apart and ground minutely wrinkled; frons with pitsbecoming smaller, closer medially, slightly distorted; at middle, a few longitudinal wrinklesand frontal line incomplete.

Mesosoma. Pronotum with short, mossy plumose hair along collar between lobes. Scutumwith distinct pits 1-2 pit widths apart, ground granular, slightly lustrous. Scutellum withpits slightly larger, ground lustrous. Mesepisternum with round pits about 1 pit widthapart, and medium-length, yellowish white, weakly plumose hair; ground uneven, roughened,

HAWAIIAN HYLAEUS 87

slightly lustrous. Hypoepimeron impunctate, smooth, lustrous; nearly flat. Basal area ofpropodeum lineately rugose to brow and beyond, shiny surface between rugae that are 2-4 rugae apart; nearly smooth below brow.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, pale,appressed hairs spaced about one hair length apart.

Color. Head and mesosoma black; legs dark brown, metasoma dark reddish brown;unmarked except fore tibia with pale area. Wings yellowish clear.

Perkins’ collections. Kauai: “Kauai” (BMNH).Recent collections. None.Flower records. None.Remarks. Colors of the integument and pubescence are based on an old specimen. This

species was not encountered in collecting at Polihale, an excellent coastal site for Hylaeuson Kauai; however, populations at such an arid site are highly cyclical, and it is possiblethat even if it occurs there, the three brief collecting trips may have missed it.


Hylaeus (Nesoprosopis) flavifrons (Kirby)(Fig. 25)

Prosopis flavifrons Kirby, 1880:85 (♂; holotype ♂ BMNH 17a 170; Kauai; wing length,3.7 mm).

Nesoprosopis flavifrons, Perkins, 1899:101 (♂♀; Pl. 2, Fig. 18).Hylaeus (Nesoprosopis) flavifrons, Meade-Waldo, 1923:34 (list).Nesoprosopis flavifrons, Daly, 1994:114 (holotype ♂ confirmed).

DIAGNOSIS. Small bees with short malar spaces and clear wings. Male face with onelarge yellow mark extended at sides to level of antennal sockets and supraclypeal area; legsunmarked; scape unmarked or with narrow, pale lateral margin; unusual scape moderatelyto strongly dilated and strongly arched. Differs from its sister species, H. anthracinus, bythe dilated S8; the facial marking also usually larger. Female black and unmarked; mandiblewith three teeth, a feature shared with H. anthracinus; distinguished from sympatric speciesby the combination of very short scutum hair and long, dark brown hair on T6.

Distribution. Kauai; coast.Male. Measurements: Malar space, 5; eye, 128:108:102:67; clypeus, 64×67; frons,

23×22:86; interalveolar, 17:26:47:58, D 18; interocellar, 28:28:15:31, D 15; scape, 41×39;wing length, 3.8 mm (Kauai, BPBM 3387).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area with punctationindistinct, pits about 1 pit width apart, partially obscured and distorted by weak parallel,minute wrinkles that curve from near the lateral ocellus to along the eye (pits more distinctand wrinkles weaker in some bees, in these the size and density of pits is rather uniformacross the frons); frons with pits medially distinct, nearly round, about 1 pit width apart;ground medially smooth, lustrous; fine frontal line evident in some bees; impunctate, lustrousarea above antennal sockets. Lower paraocular depression very shallow. Frontoclypealsulcus slightly arched. Scape in frontal view moderately to strongly dilated, width about


equal to length, ratio about 0.8-1.0; lateral edge in frontal view distinctly convex; stronglyarched in lateral view with maximum arch near middle of scape; no groove on underside;with short to medium-length, white, plumose hair. Vertex hair 12-20, yellowish white,plumose.

Mesosoma. Scutum with distinct pits 1-2 pit widths apart, ground nearly smooth, slightlylustrous; hair short (about 6), white, plumose. Scutellum with pits slightly larger, groundsmooth, lustrous. Mesepisternum with distinct, round pits about 1 pit width apart, andlong, white, plumose hair; ground nearly smooth, lustrous. Hypoepimeron impunctate,smooth to weakly striate, lustrous; slightly swollen. Metanotum slightly lustrous. Basalarea of propodeum irregularly lineate or reticulately rugose to brow; shiny between rugae;brow minutely wrinkled, slightly lustrous.

0.5 mm

A B C D

E F

1.0 mm

Fig. 25. Hylaeus flavifrons, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

HAWAIIAN HYLAEUS 89

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, pale,relatively long, appressed hairs spaced about one hair length apart or less. T7 hair short,dark brown, prostrate.

Terminalia. Lateral process of S7 parallel-sided, length greater than width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with medium-length, narrow,straplike tips. Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face with one large mark withfine black sulci in “H” shape: clypeus entirely yellow or nearly so; lower paraocular areaentirely yellow or nearly so to level of antennal socket, sometimes reduced to a broadstripe (c.f. H. anthracinus) or upper margin of mark at sides diffuse yellowish brown;supraclypeal area to acme yellow; scape unmarked or with narrow, pale lateral margin.Pronotum unmarked. Legs dark brown to black except fore tibia with pale area; unmarked.Wings clear.

Female. Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area andfrons with indistinct, shallow pits about 1 pit width apart; at sides weak parallel, minutewrinkles curved from near the lateral ocellus to along the eye, otherwise ground minutelywrinkled, slightly lustrous. Apex of clypeus shallowly emarginate. Mandible with threeteeth (see Fig. 9B). Dorsal end of facial fovea extended to level of lateral ocellus. Vertexhair 8-16, yellowish white, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with indistinct pits 1-2 pitwidths apart, ground nearly smooth, slightly lustrous; hair short (4-6), pale, weakly plumose.Mesepisternum with shallow, round pits about 1 pit width apart, and short, white, plumosehair; ground nearly smooth, lustrous. Hypoepimeron impunctate, smooth, lustrous; nearlyflat. Metanotum slightly lustrous. Basal area of propodeum irregularly lineately rugose tobrow; shiny between rugae; brow minutely wrinkled, slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, pale, short,appressed hairs spaced about one hair length apart. T6 hair long, dark brown, prostrate.

Color. Head and mesosoma dark brown to black; legs and metasoma dark brown;unmarked except fore tibia with obscure pale area. Wings clear.

Perkins’ collections. Kauai: “Kauai” (BPBM), coast (BPBM); Waimea (BPBM,OUMNH), 2000 ft (BMNH).

Recent collections. Kauai: Polihale State Park, coast. Lehua: West Horn, coast.Flower records. Portulaca oleraceae*, Scaevola sericea, Sida.Remarks. Occurs in company with H. chlorostictus, H. connectens, H. hostilis, and H.

solaris at Polihale beach on Kauai. Males are easily separated by the strongly dilated scapeand the facial marks, in which the paraocular marks are broad and usually reach the eye butdo not extend above the antennal sockets; females have very short scutum hair like H.solaris, but the hairs of T6 are dark rather than pale. A male collected from Lehua, an isletnear Niihau, has reduced facial marks similar to H. anthracinus and is almost certainlyresponsible for the report of that species from Niihau by Beardsley and Tuthill (1959).

References. Blackburn and Cameron, 1886:144, 148 (♂; key); 1887:203-204, 208 (♂;key). Dalla Torre, 1896:23 (list). Perkins, 1910:603 (key); 1913:64, 81 (biology,morphology).


Hylaeus (Nesoprosopis) flavipes (F. Smith)(Fig. 26)

Prosopis flavipes F. Smith, 1853:23 (♂; holotype ♂ BMNH 17a 153; Sandwich Islands;wing length, 4.0 mm).

Nesoprosopis flavipes, Perkins, 1899:99 (quotes Smith’s description).Hylaeus (Nesoprosopis) flavipes, Meade-Waldo, 1923:34 (list).Nesoprosopis flavipes, Daly, 1994:120 (holotype ♂ confirmed).Prosopis blackburni F. Smith, 1879a:682; 1879b:20 (♂♀) (new synonymy).Nesoprosopis blackburni, Perkins, 1899:97 (♂♀; Pl. 2, Fig. 16).Hylaeus (Nesoprosopis) blackburni, Meade-Waldo, 1923:34 (list).Prosopis blackburni, Daly, 1994:119 (lectotype ♂ BMNH 17a 2827ab; Maui, wing length,

3.7 mm).

DIAGNOSIS. Small to medium-sized bees with very short to short malar spaces and clearto slightly smoky wings. Male lower face entirely yellow, extended at sides in a broadstripe above antennal sockets, scape black or with an apical yellow spot or a yellow stripeextended the full length of the scape; tibiae and tarsi of all legs yellow with some subapicalbrown or orange spots on the tibiae visible in caudal view. Female black and usually unmarked(rare specimens may have small yellow marks in paraocular areas).

Distribution. Hawaii, Maui, Lanai (new record), and possibly Oahu; coast, and lowlandand montane dry shrubland and forest. No recent collections from Maui.

Male. Measurements: Malar space, 5; eye, 137:108:100:74; clypeus, 66×69; frons,25×24:93; interalveolar, 12:22:54:64, D 17; interocellar, 29:27:25:32, D 15; scape, 49×22;wing length, 4.3 mm (Hawaii, Hawaii Volcanoes N. P., Kalu, coast, 100 ft).

Head. Malar space very short to short (ratio about 0.2-0.3 DMO). Upper paraoculararea with round pits about 1 pit width apart or more, frons with pits becoming closer andsmaller medially; pits minute at middle, contiguous and slightly distorted; ground at sideswith weak parallel wrinkles curved from near lateral ocellus to along the eye, groundsmoother medially, lustrous; middle without special sculpture, no frontal line evident;impunctate, lustrous area above antennal sockets, extended medially to middle of frons insome bees. Lower paraocular depression very shallow to absent. Frontoclypeal sulcusstraight or nearly so. Scape in frontal view weakly to slightly dilated, width half or less oflength, ratio about 0.4-0.5; lateral edge in frontal view nearly straight or slightly concave;moderately arched in lateral view with maximum arch near middle of scape; no groove onunderside; with short to medium-length, yellowish white, weakly plumose hair. Vertex hair10-14, yellowish white, plumose.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground nearly smoothly coriaceous,lustrous; hair medium-length (8-10), yellowish white, plumose. Scutellum with pits slightlylarger, ground smoothly coriaceous, lustrous. Mesepisternum with shallow, indistinct pitsabout 2 pit widths apart, and long, yellowish white, plumose hair; ground smooth, lustrous.Hypoepimeron impunctate, smooth, lustrous; nearly flat. Metanotum slightly lustrous. Basalarea of propodeum lineately rugose, rugae reaching from the base to between a quarter

HAWAIIAN HYLAEUS 91

and almost all the way to brow, some bees with only a few widely spaced rugae or sparselyreticulate, shiny between rugae; brow minutely wrinkled, dull.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, short, pale,suberect to appressed hairs spaced about one hair length apart. T7 hair short, golden,appressed.

Terminalia. Lateral process of S7 tapered, pale, length subequal to width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with moderately long, narrow,rounded tips set at an angle to each other. Apices of gonoforceps subequal to penis valves.

Color. Labrum yellow; mandible black or partly yellow. Face with one large mark withfine black sulci in “H” shape: clypeus entirely yellow; paraocular area yellow to well above

0.5 mm

E F

A B C D

1.0 mm

Fig. 26. Hylaeus flavipes, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.


antennal socket; supraclypeal area to acme yellow. Scape in frontal view variable: entirelyblack or with an apical yellow spot, or a yellow stripe extended up to full length of scape.Flagellum orange to orange-brown beneath. With or without marks on pronotal collar(transverse collar stripe of well-separated, narrow, irregular lateral marks), pronotal lobes,and tegula. Femora brown to dark brown except marked yellow at very apex; tibiae yellowexcept in frontal view, usually with an obscure orange to brownish stain near midlength,and in caudal view with a large subapical orange to dark brown spot (especially visible onthe mid and hind tibiae); basitarsi entirely yellow, basal mediotarsus yellow, and distalterminal tarsal segments yellowish brown. Wings clear.

Female. Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular areawith distinct, round pits about 1 pit width apart or more, frons with pits closer medially,some contiguous near middle, but still distinct and mostly round; ground smooth, lustrous,especially at each side near the center of the frons; at middle, a few longitudinal wrinklesand a faint frontal line evident in some bees; a small, impunctate, lustrous area aboveantennal sockets. Apex of clypeus shallowly emarginate. Dorsal end of facial fovea extendedto level of lateral ocellus. Vertex hair 6-10, yellowish white, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground nearly smoothly coriaceous, slightly lustrous; hair short (4-6; specimensfrom montane areas on Hawaii may also have scattered long hairs up to 20), white, plumose.Scutellum with pits slightly larger, ground smoothly coriaceous, lustrous. Mesepisternumwith shallow, indistinct pits about 1 pit width apart or more, and medium-length, yellowishwhite, plumose hair; ground smooth or very weakly striate, lustrous. Hypoepimeronimpunctate, smooth, lustrous; nearly flat. Metanotum slightly lustrous. Basal area ofpropodeum lineately rugose to brow or nearly to brow, shiny between rugae; brow minutelywrinkled to nearly smooth, dull to slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, very short,pale, appressed hairs spaced more than a hair length apart. T6 hair short, light to goldenbrown, appressed.

Color. Head black, usually unmarked (rarely with small yellow spots in paraocular areas);mesosoma black; legs and metasoma dark brown; unmarked except fore tibia with palearea and with or without an obscure dorsal yellowish stripe. Wings clear to slightly smoky.

Perkins’ collections. H. blackburni: Maui: Maui coast (BPBM); Iao Valley, far up(BPBM); Wailuku (BMNH, BPBM, OUMNH), sand hills (BPBM).

Recent collections. Hawaii: Kau Desert, 300 ft; Kalu, coast; Kaulana Bay, coast; SouthPoint, coast; Puu Pa, 2600 ft; Kipuka Nene, 2900 ft; Kipuka Alala, 5800 ft; Hale Pohaku,9000 ft. Lanai: Kahue, 1400 ft.

Flower records. Dodonaea, Jacquemontia, Myoporum, Scaevola sericea, Sesbaniatomentosa, Sida, Sophora, Styphelia, Tournefortia*, Tribulus cistoides.

Remarks. Hylaeus flavipes from Hawaii and H. blackburni from Maui and Lanai aretreated here as one species under the senior name, H. flavipes. Male bees referred to thesetwo species are similar in the extensive yellow marking of the face, yellow tibiae and tarsiof all legs, sculpture of the propodeum, and genitalia. In the past the males have beendistinguished primarily by the color of the scape and mandible: black and unmarked in H.

HAWAIIAN HYLAEUS 93

flavipes, marked with yellow in H. blackburni. Recent collections reveal variation in thedegree of melanism of male bees that largely invalidates this distinction. On Hawaii (SouthPoint) the darkest males have the scape, mandible, and pronotal collar black; no yellowmark on the pronotal lobes; and the hind tibia marked with a small brown spot in front anda larger dark brown spot behind. The least melanic males have the scape with an apicalyellow stripe extended up to half the length of the scape; yellowish areas on the mandible;yellow marks on the pronotal collar and lobe; and the hind tibia with a reddish spot in frontand a smaller brown spot behind. Strict use of Perkins’ key (1910:602) would identify bothH. flavipes and H. blackburni from this one population. On Lanai and Maui, males are lessmelanic than those on Hawaii. Here males have a yellow stripe extended the entire lengthof the scape, the mandible partly or mostly yellow, pronotal lobes yellow, pronotal collarwith or without a yellow spot, and hind tibia with an orange spot in front and an orangespot or brown spot behind; they are also usually smaller. Although some differentiation incolor patterns exists between the populations on Hawaii and those on Maui and Lanai, wechose to treat the differences as geographic variation within one species.

Confusion has surrounded the taxonomy of these species. Hylaeus flavipes was describedby Smith (1853) from one male labeled “Sandw. I. Beechey” and was stated to have ablack scape. The male holotype is in The Natural History Museum, London, and matchesthe description. The collection was presumed to be from Oahu because the ship’s naturalistwas said to have prevented by illness from pursuing his research among the islands (seeIntroduction), yet we have seen no other specimens taken on Oahu. At least one otherspecies collected by the Beechey expedition, Odynerus obscurepunctatus, was collectedby Perkins only on Hawaii (Perkins 1899:60). Perkins (1899:99) assumed H. flavipes wasfrom Oahu and did not mention other specimens from Oahu or elsewhere. Later, he recordedH. flavipes from the island of Hawaii and regarded the species as distinct from H. blackburni(1910:605). Fullaway (1918:396-397) offered a contrary opinion, stating without explanation“Flavipes=blackburni.” He also recorded collections from Hawaii and described the maleas often having a yellow spot on the scape. Although the type locality remains in question,the name H. flavipes has been associated with males from the island of Hawaii that havereduced or no yellow marking on the scape.

Hylaeus blackburni was described by Smith (1879a) from one male and one femalelabeled “Maui.” The male was stated to have the scape with a yellow line in front. The malelectotype is in London and matches the description. However, the type was somehowmislabeled as Prosopis simillima, an error corrected by Houston (1981:27). Blackburn(1886:141-142) supplemented the description of H. blackburni and described considerablevariation. Perkins (1899:97) identified bees in his own collection from Maui as H. blackburniand redescribed both sexes, stating that the male had the front half of the scape yellow. Hecommented that Blackburn must have had more than one species when he wrote aboutvariation in H. blackburni.

Both H. flavipes and H. ombrias are typically coastal or lowland species that haveestablished disjunct populations above 5000 ft in the Pohakuloa area of Hawaii. They mayhave formerly been continuous with populations on the Kona coast that are now extirpated.Individuals of both species from montane areas often have longer vertex and scutum hairs


than those from the lowlands. Males of H. flavipes from upland areas may have theparaocular marks in a narrowing stripe as in H. niloticus rather than the typical broadstripe, but can easily be distinguished from that species by the extensive leg markings aswell as the dilated process of S8.

References. Blackburn and Cameron, 1886:149 (♂; key); 1887:209 (♂). Dalla Torre,1896:23 (list). Perkins, 1910:602, 605 (key, record: Hawaii, coast). Fullaway, 1918:396,397 (♂♀; record: Hawaii, Honuapo, Hilea, Kau). Hylaeus blackburni: Blackburn andKirby, 1880:85 (record). Blackburn and Cameron, 1886:139, 141-142, 148 (♂♀; key);1887:201-202, 208 (♂♀; key). Dalla Torre, 1896:16 (list). Perkins, 1899:104 (host ofcleptoparasite: N. hilaris); 1910:602 (key); 1913:79 (biogeography). Fullaway, 1918:396(record: Lanai, coast).

Hylaeus (Nesoprosopis) fuscipennis (F. Smith)(Fig. 27)

Prosopis fuscipennis F. Smith, 1879a:682-683; 1879b:20-21 (♂♀; holotype ♂ BMNH17a 155; Oahu; wing length, 7.7 mm).

Nesoprosopis fuscipennis, Perkins, 1899:106-107 (♂♀).Hylaeus (Nesoprosopis) fuscipennis, Meade-Waldo, 1923:34 (list).Nesoprosopis fuscipennis, Daly, 1994:120 (holotype ♂ confirmed).Nesoprosopis fuscipennis obscuripes Perkins, 1899:107 (♂♀) (new synonymy).Hylaeus (Nesoprosopis) fuscipennis obscuripes, Meade-Waldo, 1923:34 (list).Nesoprosopis fuscipennis obscuripes, Daly, 1994:117 (lectotype ♂ BMNH 17a 140;

Maui, West Maui Mts., Iao Valley; wing length, 7.1 mm).Nesoprosopis fuscipennis swezeyi Cockerell, 1926:308 (♂♀; Oahu, Mt. Tantalus) (new

synonymy).Nesoprosopis fuscipennis swezeyi, Daly, 1994:114 (types not located).Nesoprosopis caeruleipennis Perkins, 1899:107 (♂♀) (new synonymy).Hylaeus (Nesoprosopis) caeruleipennis, Meade-Waldo, 1923:34 (list).Nesoprosopis caeruleipennis, Daly, 1994:116 (lectotype ♂ BMNH 17a 138a; Molokai,

Mts.; wing length, 7.9 mm).

DIAGNOSIS. Large bees with punctured terga, dark wings, and dark faces. Males withyellowish stripe on scape; both sexes with a complete or nearly complete transverse stripeon the pronotal collar, and no hump on second sternum as seen in lateral view. Sisterspecies of H. pubescens, which has a hump on S2 and much more, redder hair on the apexof the metasoma.

Distribution. Maui, Lanai, Molokai, and Oahu; montane wet and mesic forest. No recentcollections from Oahu.

Male. Measurements: Malar space, 6; eye, 179:146:151:102; clypeus, 97×90; frons,33×44:113; interalveolar, 20:36:56:67, D 27; interocellar, 36:44:33:44, D 18; scape, 61×51;wing length, 7.5 mm (Maui, West Maui Mtns, BPBM 3402).

HAWAIIAN HYLAEUS 95

Head. Malar space short (ratio about 0.3-0.4 DMO). Upper paraocular area withindistinct, shallow, somewhat distorted pits about 1 pit width apart, in some bees more orless separated by fine parallel wrinkles that curve across the sides of the frons from near thelateral ocellus to along the eye; frons with pits near the center relatively distinct, nearlyround, and with diminished wrinkles; ground generally slightly lustrous, more lustrousnear center; middle with some fine longitudinal wrinkles, with or without a fine, frontalline; no impunctate area above antennal sockets. Lower paraocular depression distinct.Frontoclypeal sulcus arched or nearly straight. Scape in frontal view moderately dilated,width more than half length, ratio about 0.8; lateral edge in frontal view variable, distinctlycurved (convex) or forming a broad obtuse angle at middle, to nearly straight; stronglyarched in lateral view with maximum arch near apex of scape; underside of median edgewith a narrow groove; with medium-length, yellowish white, mostly simple hair. Vertexhair 12-20, white, mostly simple.

Mesosoma. Scutum with distinct pits 1-2 pit widths apart, ground slightly lustrous; hairmedium-length (10-12), yellowish white, mostly simple. Scutellum with pits slightly larger,lustrous. Mesepisternum with distinct, round pits about 1 pit width apart, and long, yellowishwhite, plumose hair; ground smoothly coriaceous, lustrous. Hypoepimeron punctured inlower part, smooth, slightly lustrous; swollen. Metanotum dull. Basal area of propodeumslightly protuberant in middle, roughened overall by reticulate rugae except the posterior,vertical, coriaceous face; rugae and ground minutely wrinkled, slightly lustrous; lateralcarina strong, terminated anterolaterally in a slightly protuberant area at each side of thearea.

Metasoma. Disk of T2 with distinct punctation, minute, shallow pits more than 2 pitwidths apart, and fine, short, pale, appressed hair; ground smoothly coriaceous, shiny. T7hair long, reddish brown, erect. T5-6 hair not particularly long and dense.


Color. Labrum and mandible blackish brown to black. Face marks absent. Scape in frontalview with yellowish ivory stripe on lateral margin. Pronotum with ivory transverse collarstripe complete or nearly so; lobes unmarked. Legs dark brown to black except: fore femurwith or without (obscure in H. fuscipennis holotype) ivory spot at apex; fore tibia withpale area and with or without a trace of ivory at base; mid tibia with or without a trace ofivory at base and subapex, mid basitarsus with some ivory marking; hind femur with orwithout apex ivory; hind tibia with ivory stripe in frontal view up to basal three-fourths ofsegment; hind basitarsus with some basal ivory marking. Wings smoky to dark brown;some bees with iridescence.

Female. Head. Malar space medium-length (ratio about 0.6 DMO). Upper paraoculararea with shallow, slightly distorted pits about 1 pit width apart and more or less separatedby fine, parallel wrinkles; frons with pits more distinct and round, and wrinkles less evidentmedially; fine longitudinal wrinkles at middle, middle from median ocellus to acme notdepressed; with or without a fine frontal line. Apex of clypeus shallowly emarginate. Apex


of mandible with three teeth. Dorsal end of facial fovea extended to level of lateral ocellus.Vertex hair 12-20, yellowish white, weakly plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground slightly granularly coriaceous, slightly lustrous; hair long (14-20), white,weakly plumose. Scutellum with pits slightly larger, ground smoothly coriaceous, lustrous.Mesepisternum with distinct, round pits about 1 pit width apart, and long, yellowish whiteor brownish white, plumose hair; ground smoothly coriaceous, lustrous. Hypoepimeronpunctured, smooth, lustrous; swollen. Metanotum dull. Basal area of propodeum slightlyprotuberant in middle, roughened overall by reticulate rugae except the posterior, vertical,coriaceous face; rugae and ground minutely wrinkled, slightly lustrous; lateral carina strong,terminated anterolaterally in a slightly protuberant area at each side of the area.

0.5 mm

A B C D

E F

1.0 mm

Fig. 27. Hylaeus fuscipennis, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

HAWAIIAN HYLAEUS 97

Metasoma. Disk of T2 with distinct punctation; minute, shallow pits more than 2 pitwidths apart, and fine, short, pale, appressed hair; ground smoothly coriaceous, shiny. T6hair long, reddish brown, erect. T4-5 hair not particularly long and dense.

Color. Body and legs black. Face and legs unmarked. Pronotum with ivory transversecollar stripe complete or nearly so; lobes unmarked. Wings smoky to dark brown; somebees with iridescence.

Perkins’ collections. Nesoprosopis fuscipennis: Oahu: Honolulu Mts., 1500 ft (BPBM,OUMNH), 1800 ft (BPBM); Koolau Range (BPBM), 1500 ft (BMNH, BPBM), 2000 ft(BMNH, BPBM); Koolau Range, N.W., 1500 ft (BPBM); Pali (OUMNH); Tantalus(BPBM); Wahiawa, 1500 ft (BPBM); Waialua, 1800 ft (BPBM); Waianae Mts, 4000 ft(BMNH); Waimea Mts (BPBM). N. fuscipennis obscuripes: Maui: Iao Valley (BMNH,BPBM, OUMNH), far up valley (BPBM); West Maui Mts. (BMNH, BPBM, OUMNH).N. caeruleipennis: Lanai: center of Lanai, forest above Koele, 2000 ft (OUMNH);Halepaakai, 3000 ft (BPBM, OUMNH); “Lanai,” 2000 ft (BMNH), 2600 ft (OUMNH);Mts. Koele, 2000 ft (BPBM, OUMNH), 2500 ft (BPBM). Molokai: “Molokai” (OUMNH),3000 ft (BPBM), 4000 ft (BPBM). Molokai Mts., 3000 ft (BMNH, BPBM), 4000 ft(BMNH, BPBM).

Recent collections. Maui: Kaulalewelewe, 3000 ft; Puu Kukui Rd., 2200 ft; Kahoma,2600 ft. Lanai: Munro Trail, 2800-3000 ft; Kaiholena Ridge, 2400 ft. Molokai: KamakouRd., 3200 ft; Puu Kolekole trail, 3600 ft.

Flower records. Acacia confusa*, Cheirodendron, Metrosideros, Psychotria mauiensis,Tetraplasandra oahuensis.

Remarks. Perkins (1899:107) described H. caeruleipennis from Molokai and Lanai asstructurally similar to H. fuscipennis from Oahu, but differing in having blue iridescentwings and dark tarsi. He also distinguished H. fuscipennis variety obscuripes from WestMaui by violet iridescent wings and dark tarsi, noting that some from Oahu are similarwhereas others lack the iridescence. Cockerell (1926) later named two males and a femalefrom Mt. Tantalus with the antennae ferruginous beneath and the tarsi dark as a newmutation, H. fuscipennis swezeyi. The populations on Oahu, Molokai, Lanai, and Maui arestructurally similar and are considered here to be one species under the senior name, H.fuscipennis, with variation in the color of wing iridescence, extent of ivory markings, anddegree of cuticular melanization. Iridescent colors are still visible on bees collected byPerkins. The distal margins of the wings of bees from all islands may exhibit faint bands ofred and green. Recently collected bees from Molokai and Lanai may have the distal marginsblue and the center of the wing with magenta reflections, especially the underside. Beesfrom Oahu and Maui may lack color reflections or have magenta or violet reflections, justas described by Perkins. Variation in wing color in the same population of the sister species,H. pubescens, was noted by Perkins in his key (1910:604).

Hylaeus pubescens of Hawaii is the sister taxon to H. fuscipennis of Oahu and MauiNui. Both sexes of both species have the propodeum similar in shape and sculpture. Maleshave a similar scape in frontal view with a broad, obtuse angle along the lateral edge andthe median process of S8 strongly dilated. The second sternum of both sexes of H.fuscipennis, seen in lateral view, are evenly convex but without a distinct hump. Hylaeus


pubescens is distinguished by the second sternum with an angular protuberance at midlength.The protuberance may be slight or well developed, especially in females.

References. Blackburn and Kirby, 1880:85 (♂♀). Blackburn and Cameron, 1886:140,148 (record, key); 1887:200, 208 (record, key). Dalla Torre, 1896:23 (list). Perkins,1907a:43 (record: Oahu, Tantalus); 1910:604 (key); 1913:79 (biology; parasitoid:Eupelmus). Bridwell, 1919:135 (morphology). Cockerell, 1926:308-309 (record: Oahu,Mt. Tantalus). Beardsley, 1976:188, 209 (parasitoid: Coelopencyrtus kaalae). Hylaeusfuscipennis obscuripes: Perkins, 1907a:43 (record: Oahu, Tantalus); 1913:81 (parasitoid:Eupelmus). Hylaeus caeruleipennis: Perkins, 1910:604 (key). Cockerell, 1926:309(comparison with N. f. swezeyi). Swezey and Bryan, 1929:295, 297, 306 (parasitoid:Nesencyrtus kaalae; nest: in rotten Metrosideros collina polymorpha; record: Molokai).Beardsley, 1976:188, 209 (parasitoid: Coelopencyrtus kaalae).

Hylaeus (Nesoprosopis) gliddenae Magnacca and Daly, new species(Fig. 28)

DIAGNOSIS. Large bees with punctured terga and slightly smoky wings. Head andmesosoma black, metasoma red. Male with unusual narrow, linear marks along eyes andapex of clypeus; mandible and scape also marked with yellow. Female unknown. Sisterspecies of H. paradoxicus, differing in the presence of face marks and other facial characters;see remarks.

Distribution. Hawaii; probably from montane mesic forest. Known only from onecollection.

Male. Measurements: Malar space, 6; eye, 160:132:136:94; clypeus, 88×86; frons,38×34:94; interalveolar, 24:30:46:60, D 26; interocellar, 34:32:28:38, D 18; scape, 56×42;wing length, 6.7 mm (holotype).

Head. Malar space short (ratio 0.3 DMO). Upper paraocular area with distinct pitsabout 1 pit width apart, ground slightly lustrous; frons with pits closer medially but nottouching, ground dull. Lower paraocular depression indistinct. Face overall very flat, clypeusnot convex longitudinally or laterally; frons also flat. Frontoclypeal sulcus slightly arched.Scape in frontal view moderately dilated, width more than half length, ratio about 0.8;lateral edge in frontal view straight, moderately arched in lateral view, expandedposteroapically so that arch of underside is much stronger than that of upper surface; nogroove on underside; with medium-length, yellowish white, mixed simple and weaklyplumose hair. Vertex hair 20-30, white, weakly plumose.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground granularly coriaceous, lustrous;hair medium-length to long (10-20), yellowish white, simple or weakly plumose. Scutellumwith slightly larger pits evenly spaced about 1 pit width apart, ground similar to scutum.Mesepisternum with shallow but distinct pits about 1 pit width apart or less, and long,yellowish white, plumose hair; ground smooth, lustrous. Hypoepimeron shallowly punctate,slightly lustrous; swollen. Metanotum dull, roughened. Basal area of propodeum protuberantin middle, roughened overall by reticulate rugae except the posterior, vertical, coriaceous

HAWAIIAN HYLAEUS 99

Fig. 28. Hylaeus gliddenae, new species, male. A. Seventh sternum, ventral view. B. Eighth sternum,ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E.Face with left scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E.

0.5 mm

1.0 mm

A B C D

E F

face; rugae and ground granular or minutely wrinkled, dull to slightly lustrous; lateralcarina weak, terminating anterolaterally in a protuberant area.

Metasoma. Disk of T2 with distinct but weak punctation, minute, shallow pits about 3pit widths apart; ground smoothly coriaceous, lustrous; with fine, pale, short hair. T7 hairlong, reddish brown, erect.

Terminalia. Lateral process of S7 broadly triangular, length less than width ofhemisternite. Median process of S8 weakly dilated but rather broad, apex bifurcate in a Yshape with long tips; the more sclerotized dorsal bar of the process extends onto thebifurcations. Apices of gonoforceps slightly longer than penis valves, inwardly pointed.

Color. Anterior third of mandible yellow. Face with a U-shaped mark: paraocular areaswith narrow stripes along the eyes, not quite reaching to antennal socket, connected by anapical mark on the clypeus. Scape in frontal view with the lateral third to half yellow.


Pronotum unmarked. Head and mesosoma black; metasoma red; T2 with an elongate blackspot laterally behind spiracle. Legs dark brown, unmarked except pale area on fore tibia;anterior yellow marks on the basal third of the mid tibia and basal half of hind tibia; and allbasitarsi with anterior yellow marks down most or all of their length. Wings slightly smoky.

Female. Unknown.Holotype ♂ labeled “Kilauea, Hawaii, 7-16-34. O. H. Swezey collector. Nest in burrow

of Holcobius in Suttonia [= Myrsine]” (BPBM).Flower records. None.Etymology. The species epithet gliddenae is in honor of Catherine Glidden, formerly of

Hawaii Volcanoes National Park, whose assistance was crucial to KM’s study of Hylaeus.Remarks. This species is unmistakable due to the red metasoma, shared only with H.

paradoxicus, and the unusual facial marks. It is obviously the sister of that species; it alsoshares the unique extension of the sclerotized portion of the process of S8 onto thebifurcations, and the nearly triangular S7. It differs in a number of facial characters: theyellow marks on the face and mandibles and larger marks on the scape; a more stronglyarched scape with the lateral edge straight rather than convex; the clypeus flat nearly to theapex rather than slightly convex ventrally; and the median ocellus much less prominent. Italso has the sclerotized part of the S8 bifurcation longer and marks on the basitarsi andhind tibiae, and is considerably smaller than H. paradoxicus. The facial marks and discoloredscape are similar to H. anomalus, and it may be an intermediate between the two. Thefemale is probably similar to the female of H. paradoxicus and may not be distinguishablefrom a small individual of that species.

Hylaeus (Nesoprosopis) haleakalae (Perkins)(Fig. 29)

Nesoprosopis haleakalae Perkins, 1899:87-88 (♂♀).Hylaeus (Nesoprosopis) haleakalae, Meade-Waldo, 1923:34 (list).Nesoprosopis haleakalae, Daly, 1994:117 (lectotype ♂ BMNH 17a 122; Maui, Haleakala,

5000 ft, wing length, 4.6 mm).Nesoprosopis neglecta Perkins, 1899:89 (♀; holotype ♀ BMNH 17a 125; Molokai,

Mts., 4500 ft; wing length, 5.8 mm) (new synonymy).Hylaeus (Nesoprosopis) neglectus, Meade-Waldo, 1923:34 (list).Nesoprosopis neglecta, Daly, 1994:118 (holotype ♀ confirmed).

DIAGNOSIS. Medium-sized bees with smoky wings and punctured terga. Pubescencelong all over body, legs mostly black with only traces, if any, of yellow marks, unusualbasal area of propodeum deeply reticulately rugose to brow (less so in female). Male withthree separate or adjacent facial marks, wide supraclypeal area, and middle of frons withblack spot of dense punctures; unusual scape not arched and with long brown hair alongmedian edge; long brown hair on vertex and scutum. Female black and marked only insome females by short yellow stripes in lower paraocular areas.

Distribution. Maui and Molokai; montane wet forest.

HAWAIIAN HYLAEUS 101

Male. Measurements. Malar space, 7; eye, 120:130:127:86; clypeus, 71×71; frons,17×37:84; interalveolar, 22:28:49:61, D 22; interocellar, 32:42:22:39, D 15; scape, 47×29;wing length, 5.2 mm (Molokai, 4000+ ft, BPBM 3515).

Head. Malar space medium (ratio about 0.5 DMO). Upper paraocular area and fronswith pits less than 1 pit width apart, crowded and distorted in subpolygonal shapes; a darkrhomboid spot at the middle created by dense dark brown hair and very dense punctation;shiny area just below spot and above antennal socket. Lower paraocular depression absent,surface nearly flat. Frontoclypeal sulcus nearly straight or arched; wide supraclypeal area.Scape in frontal view slightly dilated, width a little more than half length, ratio about 0.6;lateral edge in frontal view nearly straight; not arched in lateral view (but arched in medianor posterior view); underside of median edge with a groove; and long, simple, pale tobrown hair. Vertex hair 20-30 or more, brown, simple.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground slightly lustrous; hair long(14-20), reddish to dark brown, mostly simple. Scutellum with pits larger. Mesepisternumwith distinct, round pits, about 1 pit width apart, and long, brownish white, weakly plumosehair; ground minutely coriaceous, slightly lustrous. Hypoepimeron impunctate, smooth,lustrous, slightly swollen. Metanotum dull. Basal area of propodeum reticulately rugose inentire anterior area and brow; areolae varying from large and deep with ground shiny tosmall and shallow with ground granular.

Metasoma. Disk of T2 with shallow pits; hair short, dark, erect, relatively dense,conspicuous in lateral view; pits and hair more conspicuous on T3-7. T6-7 hair long, black,erect.

Terminalia. Lateral process of S7 parallel-sided, length greater than width of hemisternite.Median process of S8 strongly dilated, apex bifurcate with broad tips. Apices of gonoforcepsexceeding penis valves.

Color. Head and mesosoma black; metasoma dark brown to black. Labrum and mandibleblackish brown to black. Face with three separate or adjacent marks: clypeal mark varyingfrom a subapical, irregular yellow spot to entirely yellow; in the latter case the supraclypealarea sometimes with an irregular yellow mark; lower paraocular area with irregular triangularyellow mark along the eye below antennal socket, or completely filled in with color extendingabove antennal socket; supraclypeal area and scape unmarked. Pronotum unmarked. Legsdark brown to black except: fore femur with pale area and yellow apex; fore tibia with palearea and with only a trace of yellow at base; mid basitarsus with or without a trace ofyellow at base; hind tibia and basitarsus with or without a trace of yellow at base. Wingssmoky.

Female. Head. Malar space short (ratio about 0.4 DMO). Face convex in lateral view.Upper paraocular area and frons convex; upper paraocular area with nearly round pits lessthan 1 pit width apart, ground lustrous to shiny; frons with pits becoming crowded mediallyand partly distorted in shape, and at the middle a few, fine, longitudinal carinulae; frontalline distinct only near acme; small shiny area just above antennal socket. Apex of clypeusdistinctly emarginate. Dorsal end of facial fovea extended to level of lateral ocellus. Vertexhair 16-30, brown, simple.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground slightly lustrous; hair long (14-20), red-brown, weakly plumose.


Mesepisternum with indistinct, round pits, about 1 pit width apart; ground minutelycoriaceous, slightly lustrous. Hypoepimeron mostly impunctate, smooth, lustrous, slightlyswollen. Metanotum dull. Basal area of propodeum reticulately rugose on anterior areaextended indistinctly to brow; rugosity less pronounced than in male.

Metasoma. Disk of T2 with some micropits at hair bases, smoothly coriaceous, lustrous,with fine, pale, appressed hairs spaced about one hair length apart; pits more distinct onT3-6. Hair of middle terga much less dense and conspicuous than in male. T6 hair long,black, strongly erect.

Color. Body and legs dark brown to black except fore tibia with pale area. Lowerparaocular area with or without an obscure yellow stripe or trace along eye; clypeus andsupraclypeal area unmarked. Pronotum unmarked. Wings smoky.

0.5 mm

A B C D

E F

1.0 mm

Fig. 29. Hylaeus haleakalae, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


Perkins’ collections. Nesoprosopis haleakalae: Maui: Haleakala, 4000 ft (BMNH,BPBM); 5000 ft (BMNH, BPBM); >5000 ft (BPBM). N. neglecta: Molokai: “Molokai,”4000 ft (BPBM, OUMNH); Molokai Mts., 4000 ft (BPBM); 4500 ft (BMNH).

Recent collections. Maui: Puu Kukui Tr., 3700 ft; Makawao Forest Reserve, 4300 ft.Molokai: West Kawela Gulch, 3600 ft; Kawela Rd., 3900 ft.

Flower records. Cheirodendron, Metrosideros, Scaevola chamissoniana.Remarks. Fullaway (1918:394) pointed out the probable synonymy of H. neglectus

from Molokai and H. haleakalae from Maui, both published simultaneously by Perkins in1899. The two names are considered here to apply to one species. The name H. haleakalaeis chosen because it is based on both sexes. Lacking a male, Perkins described H. neglectusfrom a single female and compared it to H. laticeps, now known as H. unicus. No mentionwas made of a relationship with H. haleakalae. It is now reasonable to conclude that H.neglectus is the female of H. haleakalae. Recent collections from the same places on Molokaiinclude males referable to H. haleakalae and females referable to H. neglectus. Males onMaui and Molokai have the distinctive coarse, clathrate (latticed) rugosity described byPerkins for the propodeum of H. haleakalae. Females on both islands have a similarpropodeal rugosity less deeply sculpted. The type female of H. neglectus and three otherfemales from Molokai are similar to females of H. haleakalae from Maui, but lack thesmall facial marks seen in some H. haleakalae. In comparing specimens from Molokai,West Maui, and East Maui, in that order, trends are apparent of increasing facial markingsin both sexes, stronger rugae with smoother areolae in the propodeum, and shorterbifurcations of the process of S8.

References. Perkins, 1910:602 (key). Swezey, 1931:502 (nest: in dead stem of Rubushawaiiensis; record: Maui, Olinda); 1954:183 (nest: dead stem of Rubus hawaiiensis; record:Maui, Olinda).

Hylaeus (Nesoprosopis) hilaris (F. Smith)(Fig. 30)

Prosopis hilaris F. Smith, 1879a:683; 1879b:21 (♂; holotype ♂ BMNH 17a 171; Maui;wing length, 4.1 mm).

Nesoprosopis hilaris, Perkins, 1899:103-104 (♂♀; cleptoparasite of N. blackburni, N.assimulans, probably N. anthracina).

Hylaeus (Nesoprosopis) hilaris, Meade-Waldo, 1923:34 (list).Nesoprosopis hilaris, Daly, 1994:120 (holotype ♂ confirmed).

DIAGNOSIS. Medium-sized cleptoparasitic bees with clear wings, red basal terga, andunusual pale hair bands on apical terga (may be indistinct or lost by abrasion). Male lowerface entirely yellow, extended at sides in a broad stripe above antennal sockets, scapemoderately dilated and pale yellow. Female paraocular areas not marked, face and antennaemore or less red, clypeus emarginate, fore tarsus with straight hair. See discussion ofcleptoparasites in remarks.

Distribution. Maui, Lanai, and Molokai; coast. No recent collections from Maui orLanai.


Male. Measurements: Malar space, 3; eye, 146:123:118:82; clypeus, 69×72; frons,28×23:97; interalveolar, 18:28:56:67, D 20; interocellar, 31:31:28:38, D 18; scape, 51×34;wing length, 4.7 mm, tip damaged (Maui, Wailuku, BPBM 3424).

Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular area with roundpits about 1 pit width apart; frons with pits becoming smaller medially, closer (some pitscontiguous) at middle, but still distinct and nearly round; ground generally smooth andlustrous across areas; no frontal line or other special sculpture at middle; impunctate, lustrousarea above antennal sockets. Lower paraocular depression absent. Frontoclypeal sulcusstraight or nearly so. Scape in frontal view moderately dilated, widest subapically, widthmore than half length, ratio about 0.6-0.7; lateral edge in frontal view nearly straight;slightly arched in lateral view with maximum arch near middle of scape; no groove onunderside; with short, white, plumose hair. Vertex hair 12-20, yellowish white, plumose.

Mesosoma. Scutum with distinct pits about 1 pit width apart, ground smooth to slightlygranular, slightly lustrous; hair short (4-8), white, plumose. Lateral margin of front coxawith very narrow carina (see Fig. 8C), not expanded or produced. Mesepisternum withshallow, indistinct pits about 1 pit width apart, and medium-length, white, plumose hair;ground nearly smooth, lustrous. Hypoepimeron impunctate, smooth, lustrous; slightlyswollen. Metanotum slightly lustrous. Basal area of propodeum short, flat, without definitebrow; sculpture variable, reticulately rugose at front and shiny between rugae, becomingweakly reticulate and dull apically; varying to sparsely lineately rugose (some bees withtransverse rugae) and shiny between rugae, nearly smooth apically.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale, simple,appressed hairs spaced about one hair length apart; T2-6 in unworn specimens withtransverse apical bands of plumose hair, forming thin white fasciae. T7 hair short, goldenyellow, prostrate.

Terminalia. Lateral process of S7 tapered, length greater than width of hemisternite.Median process of S8 weakly dilated, apex bifurcate, tips medium-length, narrow, andstraplike. Apices of gonoforceps subequal to penis valves.

Color. Labrum yellowish brown to brown; mandible yellow at base. Face with one largemark divided by fine black sulci: clypeus entirely yellow or nearly so, supraclypeal area toacme yellow, paraocular area entirely yellow, with a broad mark extended above antennalsocket; scape in frontal view mostly yellow, flagellum yellow-brown; some bees withadditional small yellowish spot below the median ocellus. Pronotum with or without yellowtransverse collar stripe at each side; lobes yellowish brown; tegulae with or without ayellow spot. Legs yellow to yellowish brown except femora and trochanters darker, brownto black; fore tibia with indistinct pale area; hind tibia with or without diffuse yellowishstripe at base, slightly darker subapically. Head and mesosoma dark brown to blackishbrown; metasoma variable: basal 1-3 segments red and apical segments dark reddish brownor reddish black, or all segments reddish brown. Wings clear.

Female. Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular areaand frons surfaces with indistinct, shallow round pits about 1 pit width apart; ground nearlysmooth and lustrous; faint frontal line at middle; impunctate, lustrous area above antennalsockets. Apex of clypeus deeply emarginate. Dorsal end of facial fovea below level oflateral ocellus.


Mesosoma. Fore tarsus with hair straight (see Fig. 8B). Scutum with distinct pits about1 pit width apart, ground nearly smooth, slightly lustrous; hair short (4-6), yellowish white,mostly simple. Lateral margin of front coxa with very narrow carina (see Fig. 8C), notexpanded or produced. Mesepisternum with distinct, round pits about 1 pit width apart,and short, white, weakly plumose hair; ground minutely wrinkled, dull. Hypoepimeronimpunctate, weakly striate, indistinctly lustrous, slightly swollen. Metanotum indistinctlylustrous. Basal area of propodeum short, flat, without definite brow; irregularly lineatelyrugose at front, shiny between rugae, becoming weakly reticulate, rough, and duller apically.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with very short, fine,pale, simple, appressed hairs spaced more than a hair length apart; T2 with a very narrow,inconspicuous, transverse apical band of somewhat thicker hair; T3-5 with distinct yellowish

0.5 mm

D

E F

1.0 mm

Fig. 30. Hylaeus hilaris, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.


white, thin apical fascia of slightly thicker, denser hair. T6 hair long, apparently simple,golden yellow, prostrate.

Color. Head and mesosoma dark brown to black; metasoma reddish brown. Face withoutdistinct marks, but clypeus and scape yellowish brown, paraocular areas reddish brown.Pronotum with lobes yellowish brown, but without distinct mark. Fore legs yellowish brown,mid legs darker yellowish brown, hind legs brown to dark brown at base; unmarked. Apicalterga with thin, apical bands of white, plumose hair. Wings clear.

Perkins’ collections. Maui: “Maui” (BPBM); sand hills (BPBM); Wailuku (BMNH,BPBM); sandy isthmus (OUMNH). Lanai: “Lanai” (BMNH, BPBM).

Recent collections. Molokai: Moomomi, coast.Flower records. None.Remarks. Colors of the integument and pubescence of the female are based on an old

specimen. Perkins (1899:106; 1913:lxxx) discovered five species in which the femaleshave short, straight hair on the fore tarsi and considered them closely allied: H. hilaris, H.hostilis, H. inquilina, H. sphecodoides, and H. volatilis. He associated the straight tarsalhair with cleptoparasitic habits. At least the basal metasomal segments of both sexes aretypically red to reddish brown in H. hilaris and H. hostilis, but the segments of H.sphecodoides and H. volatilis may be reddish brown or entirely black in melanic individuals(see remarks for H. volatilis). The metasoma of H. inquilina is always black. Red on theterga is not unique to this group; some unrelated nonparasitic species, such as H. paradoxicusor H. anomalus, also have red terga. Curiously, red terga have evolved repeatedly in differentlineages of other cleptoparasitic bees (Michener 2000:31). Females of all the Hawaiiancleptoparasites have the general area of the frons and vertex slightly swollen or convex.With the exception of H. inquilina, female cleptoparasites have a distinctly emarginateclypeus, the facial foveae do not or just barely reach the level of the lateral ocelli, and thefore coxae have a lamellate carina (Fig. 8D; H. hilaris also lacks the lamellate carina). Allare usually found in dry habitats – H. hilaris is restricted to the coast and H. inquilina tomontane areas, while the remaining three species may be found at nearly all elevations –and appear to parasitize only ground-nesting species. Hylaeus chlorostictus, H. difficilis,H. hirsutulus, H. laetus, H. nivicola, and H. volcanicus are probably the primary hosts inthe mountains, and H. assimulans, H. chlorostictus, H. flavifrons, H. flavipes, and H.longiceps serve as hosts at the coast.

References. Blackburn and Kirby, 1880:85 (record). Blackburn and Cameron, 1886:147-148 (♀; key); 1887:207-208 (♀; key). Dalla Torre, 1896:24 (list). Perkins, 1910:604 (key);1913:79, 81 (biogeography; cleptoparasite of N. assimulans, N. longiceps, and N.blackburni). Fullaway, 1918:396 (record: Molokai).

Hylaeus (Nesoprosopis) hirsutulus (Perkins)(Fig. 31)

Nesoprosopis hirsutula Perkins, 1899:79-80 (♂♀; Pl. 2, Fig. 10).Hylaeus (Nesoprosopis) hirsutulus, Meade-Waldo, 1923:33 (list).Nesoprosopis hirsutula, Daly, 1994:117 (lectotype ♂ BMNH 17a-158b; Kauai, Halemanu,



DIAGNOSIS. Medium-sized bees with dark smoky wings. Male face with an ivory oryellow mark (clypeus and supraclypeal area, with or without lower paraocular areas), unusuallateral process of S7 very short, median process of S8 not dilated, apex simple; unusualgonoforceps expanded apically beyond penis valves. Female black and unmarked. Sisterspecies of H. perkinsianus; see treatment of that species.

Distribution. Kauai; widespread, from dry shrubland to wet forest, mountains andlowlands, but not common.

Male. Measurements: Malar space, 3; eye, 138:133:110:81; clypeus, 68×74; frons,23×36:95; interalveolar, 20:28:60:67, D 23; interocellar, 31:41:18:31, D 17; scape, 54×25;wing length, 5.3 mm (Kauai, Halemanu, 4000 ft, BPBM 3430).

Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular area and fronswith distinct, round, shallow pits less than 1 pit width apart; ground minutely wrinkled,dull; median, impunctate, lustrous area just above antennal sockets; faint frontal line evident.Lower paraocular depression distinct, but shallow. Frontoclypeal sulcus straight. Scape infrontal view slightly dilated, width half length, ratio about 0.5; lateral edge in frontal viewslightly concave; moderately arched in lateral view with maximum arch near middle ofscape; no groove on underside; with long, pale brown, mostly simple hair along the medianedge. Vertex hair 12-22, brown, weakly plumose.

Mesosoma. Scutum with pits about 2 pit widths apart, ground rather smoothly coriaceous,slightly lustrous; hair long (approximately 14), brownish white, weakly plumose. Scutellumwith pits slightly larger. Mesepisternum with indistinct, round pits, about 1 pit width apart,and long, yellowish white, weakly plumose hair; ground minutely coriaceous, slightlylustrous. Hypoepimeron mostly impunctate, smooth, slightly lustrous, slightly swollen.Metanotum dull. Basal area of propodeum lineately rugose to brow, shiny between rugaeat front; brow granular to smoothly coriaceous, dull to slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, pale,appressed hairs spaced about one hair length apart. T7 hair short to medium, black, prostrate.

Terminalia. Lateral process of S7 very short, blunt, length less than width of hemisternite.Median process of S8 linear, apex simple or nearly so. Apices of gonoforceps exceedingpenis valves, with very long curled hairs.

Color. Labrum and mandible blackish brown to black. Face with one large mark: clypeusentirely yellow or with narrow apical margin black; lower paraocular area variable, with orwithout a narrow yellow stripe along sulci, supraclypeal area to acme yellow; scapeunmarked. Pronotum unmarked. Legs dark brown to black except: fore tibia with pale areaand basal yellow spot or short stripe; mid tibia with or without a spot of yellow at base;hind tibia with basal yellow stripe less than half length of segment. Wings dark smoky.

Female. Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular areaand frons with round, shallow pits about 1 pit width apart or less, slightly closer toward themiddle; ground smooth or minutely wrinkled, slightly lustrous; fine frontal line evident.Apex of clypeus shallowly emarginate. Dorsal end of facial fovea extended to level oflateral ocellus. Vertex hair 20-30, brown, weakly plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with pits about 2 pit widthsapart, ground minutely granularly coriaceous, slightly lustrous; hair medium-length (8-16), white, weakly plumose. Scutellum with pits slightly larger, ground smoothly coriaceous,


lustrous. Mesepisternum with distinct, round pits, about 1 pit width apart, and long, yellowishwhite, weakly plumose hair; ground minutely coriaceous to striate, slightly lustrous.Hypoepimeron mostly impunctate, smooth, slightly lustrous, slightly swollen. Metanotumdull. Basal area of propodeum lineately rugose to brow, shiny between rugae at front;brow smoothly coriaceous, slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, pale,appressed hairs spaced about one hair length apart. T6 hair short-plumose, long, brown,prostrate.

Color. Head and mesosoma black; legs and metasoma very dark brown to black;unmarked. Wings dark smoky.

0.5 mm

A B C D

E F

1.0 mm

Fig. 31. Hylaeus hirsutulus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with left scape. F. Scape,lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shown by dashed lines inB-E.


Perkins’ collections. Kauai: Halemanu, 4000 ft (BMNH, BPBM, OUMNH), 4000 ft(field note #501) (OUMNH); “Kauai,” 4000 ft (BPBM); Koholuamano (field note #522),4000 ft (BMNH, BPBM, OUMNH); Makaweli, 2000 ft (BMNH); Waimea, 4000 ft (BPBM).

Recent collections. Kauai: Alakai Wilderness, 4600 ft; Mohihi Trail, 3900 ft; NualoloCliff Trail, 2800 ft.

Flower records. Chamaesyce atrococca, Metrosideros, Styphelia.Remarks. The measured male of H. hirsutulus has an extreme ratio ”interocellar distance/

diameter of median ocellus” at 1.5 and a very high ratio “ocellus-eye distance/interocellardistance” at 1.3. The ratios indicate that the lateral ocelli are unusually close together withrespect to the diameter of the median ocellus and are well separated from the eyes.


Hylaeus (Nesoprosopis) hostilis (Perkins)(Fig. 32)

Nesoprosopis hostilis Perkins, 1899:104-105 (♂♀; cleptoparasite of N. laeta; Pl. 2,Figs. 4, 20).

Hylaeus (Nesoprosopis) hostilis, Meade-Waldo, 1923:34 (list).Nesoprosopis hostilis, Daly, 1994:117 (lectotype ♂ BMNH 17a 167b; Kauai, Waimea,

3000 ft).

DIAGNOSIS. Small cleptoparasitic bees with slightly smoky wings and red basal terga.Male lower face entirely yellow, extended at sides in a broad stripe above antennal sockets,scape slightly dilated and in frontal view yellowish on lateral margin. Female clypeusemarginate, fore tarsus with straight hair, paraocular areas with triangular ivory marks.

Distribution. Kauai; coast, montane dry shrubland and mesic forest.Male. Measurements: Malar space, 2; eye, 114:98:92:63; clypeus, 53×55; frons,

18×20:78; interalveolar, 10:24:45:57, D 20; interocellar, 24:27:22:27, D 14; scape, 39×25;wing length, 3.8 mm (H. hostilis lectotype).

Head. Malar space very short (ratio about 0.1 DMO). Upper paraocular area withindistinct pits about 1 pit width apart; frons with pits becoming distinct medially, smaller,closer, ground smooth and lustrous, at middle with dense, minute pits, dull; indistinct frontalline evident; small impunctate area above antennal sockets. Lower paraocular depressionabsent. Frontoclypeal sulcus straight or nearly so. Scape in frontal view slightly dilated,width a little more than half length, ratio about 0.6; lateral edge in frontal view nearlystraight; slightly arched in lateral view with maximum arch near base of scape; no grooveon underside; with very short, white, weakly plumose hair. Vertex hair 4-10, white, plumose.

Mesosoma. Scutum with distinct pits about 1 pit width apart, ground smooth, lustrous;hair very short (about 4), white, plumose. Lateral margin of front coxa with carina lamellate(see Fig. 8D; sometimes weakly developed). Mesepisternum with indistinct, shallow pitsabout 1 pit width apart, and medium-length, white, plumose hair; ground smooth, lustrous.Hypoepimeron impunctate, smooth, lustrous; slightly swollen. Metanotum slightly lustrous.Basal area of propodeum irregularly lineately rugose to reticulate at front, becoming morereticulate near brow, shiny between rugae at front; brow weakly reticulately rugose, dull.


Metasoma. Disk of T2 impunctate; smoothly coriaceous, lustrous; with short, pale,appressed hairs spaced about one hair length apart or more. T7 hair short, pale brown,appressed.

Terminalia. Lateral process of S7 tapered, length greater than width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with short, narrow, acute tips.Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible yellow to brown. Face with a large mark, with fine blacksulci in an “H” shape, and conspicuous tentorial pits: clypeus entirely yellow or nearly so;paraocular area yellow, extended in a broad stripe well above antennal socket; supraclypealarea to acme yellow. Scape in frontal view yellowish on lateral margin. Pronotum with

0.5 mm

A B C D

E F

1.0 mm

Fig. 32. Hylaeus hostilis, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.


lobes obscurely yellowish brown. Head black, with or without a yellow spot just behindthe eye above the mandible; mesosoma black, with or without scattered irregular red areas,especially around pronotal lobes, at base of pronotum, and around base of hind leg; tegula,humeral plate, and base of costa clear yellow. Two or three basal segments of metasomared; apical segments dark reddish brown. Legs dark brown except fore tibia with pale area,fore tarsus yellow to brown, all tibiae with obscure yellowish brown basal marks and allfemora with similar apical marks. Wings slightly smoky.

Female. Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular areaand frons with distinct, shallow, round pits about 1 pit width apart, ground smooth tominutely wrinkled, lustrous to dull; minute, longitudinal wrinkles at middle; faint frontalline evident. Apex of clypeus deeply emarginate. Dorsal end of facial foveae not extendedto level of lateral ocelli. Vertex hair 4-10, white, plumose.

Mesosoma. Fore tarsus with hair short, straight (see Fig. 8B). Scutum with distinct(indistinct in some bees) pits about 1 pit width apart, ground nearly smooth, lustrous; hairshort (4-6), white, plumose. Lateral margin of front coxa with carina lamellate (see Fig.8D). Mesepisternum with indistinct, round pits about 1 pit width apart, and short, yellowishwhite, weakly plumose hair; ground nearly smooth, lustrous. Hypoepimeron impunctate,nearly smooth to weakly striate, lustrous; slightly swollen. Metanotum slightly lustrous.Basal area of propodeum irregularly lineately rugose at front, becoming reticulate nearbrow, shiny between rugae; brow weakly reticulately rugose, slightly lustrous.

Metasoma. Disk of T2 impunctate; smoothly coriaceous, shiny; some bees largely bareor with very short, pale, appressed hairs spaced more than a hair length apart. T6 hairreddish or light brownish white, medium-length, prostrate.

Color. Head and mesosoma with various parts red, reddish brown, or dark brown;metasoma with T1-3 entirely orange, the remainder red to reddish brown. Labrum andmandibles reddish or yellowish brown. Face marks somewhat similar to those of male, butconspicuous yellow color confined to paraocular areas: clypeus and supraclypeal areabrownish to reddish yellow; paraocular areas with irregular, narrowing yellow stripe toabove antennal sockets; scape yellowish brown in frontal view. Pronotum with collar andlobes reddish or yellowish brown. Propodeum reddish except basal area below brow andlateral face. Tegula, humeral plate, and base of costa clear yellow. Legs brown to darkbrown except: fore femur at apex and segments distad yellowish brown; mid and hindtibiae with obscure basal reddish or yellowish marks; mid and hind femora red apically; alltrochanters red ventrally. Wings slightly smoky.

Perkins’ collections. ?Hawaii: Kilauea (4174, BPBM). Kauai: Halemanu, 4000 ft(BPBM); Kauai, 4000 ft (OUMNH); Makaweli, 2-3000 ft (BPBM); Waialeale,Koholuamano (=Kaholuamano) (field note #522) (BPBM); Waimea, 3000 ft (BMNH,BPBM); 4000 ft (BMNH, BPBM).

Recent collections. Kauai: Polihale State Park, coast; Nualolo Cliff Trail, 2800 ft;Awaawapuhi Trail, 3400 ft.

Flower records. Scaevola sericea, Styphelia.Remarks. See discussion of cleptoparasites under H. hilaris.References. Perkins, 1910:604 (key). Hopper and Daly, 1995:35 (record: coastal Kauai).


Hylaeus (Nesoprosopis) hula (Perkins)(Fig. 33)

Nesoprosopis hula Perkins, 1911:721-722 (♂♀; Hawaii, Kilauea; flower: Myoporum).Hylaeus (Nesoprosopis) hula, Meade-Waldo, 1923:35 (list).Nesoprosopis hula, Daly, 1994:116 (types not located).

DIAGNOSIS. Medium-sized bees with clear to slightly smoky wings. Male with unusualface rather flat, yellow clypeus and lower paraocular areas, unusual yellow stripe alonglateral edge of scape; pronotum and legs variably marked. Female with narrow, irregularyellow stripe along the eye up to level of antennal sockets; pronotal lobes and sometimescollar marked; supraclypeal area relatively long, about twice as wide as long.

Distribution. Hawaii; montane mesic forest.Male. Measurements: Malar space, 10; eye, 113:105:97:68; clypeus, 64×59; frons,

23×24:79; interalveolar, 18:20:44:51, D 19; interocellar, 29:29:18:26, D 14; scape, 39×27;wing length, 4.1 mm (Hawaii, Kilauea, 4000 ft, BPBM 3442).

Head. Malar space medium-length (ratio about 0.7 DMO). Upper paraocular area withmostly round pits less than 1 pit width apart, partly obscured by minutely wrinkled, slightlylustrous ground; frons with pits becoming smaller and closer medially. Lower paraoculardepression very shallow. Frontoclypeal sulcus arched; lateral epistomal sulcus with a distinctbend laterad at the tentorial pits. Scape in frontal view moderately dilated, width more thanhalf length, ratio about 0.7; lateral edge in frontal view nearly straight; moderately archedin lateral view with maximum arch near middle of scape; no groove on underside; withlong, yellowish white, mostly simple hair mainly along median edge. Vertex hair 12-20,pale brown, mixed simple and weakly plumose.

Mesosoma. Scutum with shallow indistinct pits 1-2 pit widths apart, ground granular,dull to slightly lustrous; hair long (12-16), pale brown, weakly plumose. Scutellum withpits slightly farther apart, slightly lustrous. Mesepisternum with barely discernible pits,about 2-3 pit widths apart, and long, yellowish white, weakly plumose hair; ground minutelycoriaceous, slightly lustrous. Hypoepimeron impunctate, smooth, lustrous, slightly swollen.Metanotum slightly lustrous. Basal area of propodeum with short basal rugae reachingfrom the base to between one-third and halfway to brow; remainder including brow smoothlycoriaceous and lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; fine, pale, semierecthair about one hair length apart. T7 hair short to medium, red-brown, erect.

Terminalia. Lateral process of S7 tapered, with length subequal to width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with rather long, straplike tips.Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face with three partly separatedmarks: clypeus mostly yellow except dark narrow margin along epistomal sulcus abovetentorial pits; lower paraocular area largely yellow below antennal socket (in some malesthe paraocular mark is partly divided longitudinally by a discolored streak); supraclypealarea with or without some yellow marking. Scape in frontal view with yellow stripe onlateral margin. Pronotum with transverse collar stripe interrupted medially or absent; lobesyellow at least in part. Fore leg dark brown to black except: tibia with pale area plus dorsal,


basal yellow stripe; basitarsus mostly yellow; basal mediotarsus yellow to light brown. Midand hind legs dark brown to black except: tibiae with basal and apical yellow marks (lattersmaller and sometimes absent on the hind leg); basitarsi mostly yellow; basal mediotarsusyellow to light brown. Wings clear.

Female. Head. Malar space long (ratio 0.8-0.9 DMO). Upper paraocular area and fronswith evenly spaced round pits 1 pit width apart or slightly less, closer only around themedian ocellus; ground coriaceous and lustrous laterally, minutely granular and slightlylustrous in center; frons somewhat swollen, bulging slightly between ocelli and antennalsockets; frontal line distinct. Apex of clypeus distinctly emarginate. Dorsal end of facialfovea extended to level of lateral ocellus. Supraclypeal area relatively long, W/L ratioabout 1.9. Vertex hair 12-20, pale brown, weakly plumose.

0.5 mm

A B C D

F

1.0 mm

EFig. 33. Hylaeus hula, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view, andapex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with left scape.F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shown bydashed lines in B-E.


Mesosoma. Fore tarsus with hairs apically curved. Scutum with shallow, indistinct pits2-3 pit widths apart, ground minutely granular, slightly lustrous; hair short to medium (6-12), yellowish white. Scutellum with pits slightly larger, lustrous. Mesepisternum withshallow, indistinct pits, 1-2 pit widths apart, and medium-length, yellowish white, plumosehair; ground minutely granular, slightly lustrous. Hypoepimeron with indistinct pits, stronglylustrous, slightly swollen. Metanotum slightly lustrous. Basal area of propodeum lineatelyrugose halfway or less to brow; brow smoothly coriaceous and lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart or more. T6 hair dark brown to black,long, erect.

Color. Body dark brown to black; unmarked. Face with paraocular stripe from tentorialpit to antennal socket. Pronotal lobes marked, with or without marks on collar. Legs brownto black except pale area on fore tibia and marks at bases of tibiae: up to basal third of fore,basal quarter of mid, and basal half of hind. Wings slightly smoky.

Perkins’ collections. None. Type collected later by Perkins at Kilauea, Hawaii.Recent collections. Hawaii: Tree Molds, 4000 ft; Puu Waawaa, 4100 ft.Flower records. Chamaesyce olowaluana, Myoporum, Santalum paniculatum.Remarks. Many collections have females of H. filicum placed with males of H. hula,

despite having not been taken together. They may have been associated based on abundance,as specimens of female H. hula and male H. filicum are much less common in thesecollections.

References. None.

Hylaeus (Nesoprosopis) inquilina (Perkins)(Fig. 34)

Nesoprosopis inquilina Perkins, 1899:102-103 (♂♀; cleptoparasite of other bee species).Hylaeus (Nesoprosopis) inquilina, Meade-Waldo, 1923:34 (list).Nesoprosopis inquilina, Daly, 1994:117 (lectotype ♂ BMNH 17a-169a; Hawaii, Kau,


DIAGNOSIS. Medium-sized cleptoparasitic bees with clear wings and body black withoutred basal terga (unlike other cleptoparasites). Male lower face almost entirely yellow,extended at sides in a broad stripe above antennal sockets, unique scape moderately dilatedand with lateral edge in frontal view distinctly concave. Female foretarsus with straighthair, clypeus shallowly emarginate (unlike other female cleptoparasites).

Distribution. Hawaii; montane dry shrubland and forest.Male. Measurements: Malar space, 6; eye, 134:118:108:82; clypeus, 63×69; frons,


Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area rather uniformlypunctured with round pits less than 1 pit width apart; frons with pits becoming contiguousmedially, ground minutely wrinkled, dull; frontal line not evident; median impunctate, lustrous


area just above antennal sockets. Lower paraocular depression absent. Frontoclypeal sulcusstraight. Scape in frontal view moderately dilated, width more than half length, ratio about0.7; lateral edge in frontal view strongly concave, apex forming subacute angle; moderatelyarched in lateral view with maximum arch nearer base of scape; no groove on underside;with medium-length, yellowish white, simple to weakly plumose hair. Vertex hair 12-24,pale brown, simple.

Mesosoma. Scutum with minute, indistinct pits about 1 pit width apart, ground minutelywrinkled, dull; hair medium-length (10-12), brownish white, weakly plumose.Mesepisternum with indistinct pits about 1 pit width apart, and short to long, white, plumosehair; ground minutely wrinkled, slightly lustrous. Hypoepimeron mostly impunctate, smooth;slightly swollen. Metanotum dull. Basal area of propodeum short, narrow, rather flat and

0.5 mm

A B C D

E F

1.0 mm

Fig. 34. Hylaeus inquilina, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.


without definite brow, roughened overall with lineate rugae at front, becoming weaker andreticulate distad, lustrous between rugae at front.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, short, pale,appressed hair, a hair length apart or more; in some bees, terga with longer, white hair atsides, creating thin fasciae (may be lost through abrasion). T7 hair very short, pale brown,appressed.

Terminalia. Lateral process of S7 tapered, pale, length subequal to width of hemisternite;hemisternites with short erect setae. Median process of S8 strongly dilated, apex bifurcatewith short, rounded tips. Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face with clypeus entirely yellowor with narrow margins black; paraocular area yellow to well above antennal socket in abroad stripe; supraclypeal area black or obscurely yellow below acme; scape unmarked.Pronotum unmarked. Legs entirely dark brown to black except fore tibia with pale area orlegs obscurely marked: fore femur with obscure apical spot, combined with yellowish dorsalstripe extended full length of segment; fore basitarsus yellow; mid tibia variable, with smallbasal and obscure apical yellow spots or with only a trace of yellow at base; hind tibiavariable, with basal yellow spot. Wings clear.

Female. Head. Malar space medium-length (ratio about 0.5 DMO). Upper paraoculararea and frons rather uniformly punctured with round pits about 1 pit width apart, groundsmooth coriaceous to minutely wrinkled, lustrous; middle with a few fine, longitudinalwrinkles, frontal line not evident. Apex of clypeus very shallowly emarginate. Dorsal endof facial foveae reaching just below or just up to level of lateral ocelli. Vertex hair 10-20,yellowish white, simple.

Mesosoma. Fore tarsus with hair short, straight (see Fig. 8B). Scutum with shallow,minute pits about 2 pit widths apart, ground nearly smoothly coriaceous, slightly lustrous;hair short (4-6), brownish white, simple. Mesepisternum with indistinct, shallow pits about2 pit widths apart, and short, yellowish white, weakly plumose hair; ground smooth tominutely wrinkled, slightly lustrous. Hypoepimeron mostly impunctate, smooth; slightlyswollen. Metanotum dull. Basal area of propodeum lineately rugose about halfway to brow,wrinkled behind, shiny between rugae at front; brow minutely wrinkled, slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, short, pale,appressed hair, spaced a hair length apart or more. T6 hair light reddish brown, medium-length, appressed.

Color. Head and mesosoma black; legs dark brown, with or without obscure marks atbases of tibiae; metasoma dark brown, unmarked. Wings clear.

Perkins’ collections. Hawaii: Kau, 4000 ft (BMNH, BPBM); Kilauea (BPBM), 4000ft (OUMNH); Kau district (OUMNH).

Recent collections. Hawaii: Kipuka Nene, 2900 ft; south of Bird Park, 1230 m; nearTree Molds, 4000 ft; Mauna Loa Rd., 2050 m; Mauna Loa, 6000 ft; Kipuka Alala, 5800 ft.

Flower records. Myoporum, Styphelia.Remarks. This species is basal among the cleptoparasites and differs from all the others

in several respects, the most obvious being the lack of any red markings (H. volatilis andH. sphecodoides may have melanic individuals). It is usually found in open scrubland on


recent lava flows, and presumably primarily parasitizes the sympatric H. difficilis and H.volcanicus. See discussion of cleptoparasites under H. hilaris.

References. Perkins, 1907b:97 (record: Hawaii, Kilauea); 1910:603 (key).

Hylaeus (Nesoprosopis) kauaiensis (Perkins)(Fig. 35)

Nesoprosopis kauaiensis Perkins, 1899:90 (♂♀).Hylaeus (Nesoprosopis) kauaiensis, Meade-Waldo, 1923:34 (list).Nesoprosopis kauaiensis, Daly, 1994:117 (lectotype ♂ BMNH 17a 162a; Kauai, Mts.

Waimea, 4000 ft; wing length, 4.5 mm).

DIAGNOSIS. Medium-sized bees with clear to smoky wings, propodeum with well-separated, lineate rugae in basal half or less, otherwise smooth to brow. Male with broadwhitish yellow mark partly divided by epistomal sulci; arched frontoclypeal suture; unusualwide supraclypeal area, W/L ratios 1.6-1.9; shiny paraocular depression; process of S8without flexure. Female marked with yellow in lower paraocular areas and at sides ofpronotal collar, pronotal lobes unmarked; closely resembling H. mutatus, but with longerhair on scutum and always lacking marks on pronotal lobes.

Distribution. Kauai; montane wet and mesic forest.Male. Measurements: Malar space, 3; eye, 120:120:115:69; clypeus, 61×69; frons,

23×38:82; interalveolar, 23:26:54:61, D 20; interocellar, 31:35:20:31, D 17; scape, 39×29;wing length, 4.5 mm (Kauai, Waimea, 2000+ ft, BPBM 3469).

Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular area and fronswith nearly round pits about 1 pit width apart, pits nearly round even near the middle,ground minutely wrinkled and slightly lustrous between pits; middle between median ocellusand acme with dense, contiguous pits, but without distinctive sculpture. Lower paraoculardepression distinct, but shallow; shiny along eye. Frontoclypeal sulcus strongly arched;supraclypeal area much wider than long, W/L ratios 1.6-1.9. Scape in frontal view moderatelydilated, width more than half length, ratio about 0.7; lateral edge in frontal view distinctlysinuous; strongly arched in lateral view with maximum arch near apex of scape; undersideof median edge with a narrow line or groove; with medium-length, brownish, simple hair.Vertex hair 16-30, brown, simple.

Mesosoma. Scutum with pits about 1 pit width apart, ground slightly lustrous; hairmedium to long (mostly 8-12, with scattered hairs up to 20), reddish brown, mostly simple.Scutellum with slightly larger pits, slightly lustrous. Mesepisternum with round, shallowpits about 1 pit width apart, ground smooth and lustrous and with long, brownish, plumosehair. Hypoepimeron with small pits, smooth, shiny; swollen. Metanotum slightly lustrous.Basal area of propodeum lineately rugose in basal half or less, spaced 2-4 rugae widthsapart; brow smoothly coriaceous and lustrous.

Metasoma. Disk of T1 with shallow, indistinct pits, smoothly coriaceous, lustrous, withfine, brown, appressed hairs spaced about one hair length apart or less. T2-7 with shallowbut distinct pits (more obvious on posterior terga); hair short, dark, erect hair, and relativelydense, conspicuous in lateral view. T7 hair short, dark brown, prostrate.


Terminalia. Lateral process of S7 tapered, with length subequal to width of hemisternite.Median process of S8 straight along dorsal edge, no flexure, strongly dilated, apex bifurcatewith short tips. Apices of gonoforceps slightly shorter than penis valves.

Color. Head and mesosoma black; metasoma dark brown. Labrum and mandible blackishbrown to black. Face with clypeus largely yellow except around edges; lower paraoculararea more or less filled in by yellow spot, usually not extended above clypeus. Supraclypealarea and scape unmarked. Pronotum with transverse collar stripe interrupted medially;lobes unmarked. Legs brown except pale area of fore tibia; unmarked. Wings slightly smoky.

Female. Head. Malar space very short (ratio about 0.1 DMO). Upper paraocular areaand frons with nearly round pits about 1 pit width apart, pits nearly round even near themiddle, ground minutely wrinkled and slightly lustrous between pits; middle between median

0.5 mm

A B C D

E F

1.0 mm

Fig. 35. Hylaeus kauaiensis, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


ocellus and acme with a fine frontal line. Apex of clypeus slightly emarginate. Dorsal endof facial fovea extended to level of lateral ocellus. Vertex hair 16-30, brown, weakly plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits, 1-2 pitwidths apart, ground dull; hair mixed, short to long (lower layer about 6, long hairs 12-20),red-brown, simple. Scutellum with slightly larger pits, lustrous. Metanotum dull.Mesepisternum with shallow pits 2-3 pit widths apart, ground smooth, lustrous; with long,yellowish white plumose hair. Hypoepimeron impunctate, nearly flat. Basal area ofpropodeum lineately rugose about halfway to brow, spaced about 2-4 ruga widths apart;brow smoothly coriaceous and lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart or more; hair brownish white to brownat apex of metasoma. T3-6 with shallow pits, less conspicuous than in male. T6 hair long,dark brown, erect.

Color. Head and mesosoma black; metasoma nearly black. Face with yellow line next toeye in lower paraocular area below the level of the antennal socket; clypeus and supraclypealarea unmarked. Pronotum with yellow transverse collar stripe interrupted medially; lobesunmarked. Legs nearly black except with pale area on fore tibia. Wings clear to smoky.

Perkins’ collections. Kauai: “Kauai,” 4000 ft (OUMNH); Waimea, >2000 ft (BPBM),3-4000 ft (May), 4000 ft (IV, VI) (BMNH); Waimea Mts., >3000 ft (BPBM).

Recent collections. Kauai: Mt. Kahili, 2500 ft; Alakai Swamp Trail, 3600 ft; KahuamaaFlat, 3800 ft; Mohihi Trail, 4000 ft.

Flower records. Metrosideros, Scaevola procera, Styphelia.Remarks. Fullaway (1918:394) pointed out the possible synonymy of H. kauaiensis

with H. unicus and H. binominatus, known then as N. laticeps. However, the process ofS8 is without a flexure and the gonoforceps are slightly shorter than the penis valves in H.kauaiensis, whereas the other two taxa, synonymized below under H. unicus, have theprocess of S8 with a flexure and the gonoforceps subequal to the penis valves. The apex ofthe metasoma of female H. kauaiensis has conspicuous dense, erect hair as in H. haleakalae.

References. Perkins, 1910:603 (key). Swezey, 1954:177 (nest: in Tetraplasandra (referredto as Pterotropia) twig; record: Kauai, Halemanu).

Hylaeus (Nesoprosopis) kokeensis Magnacca and Daly, new species(Fig. 36)

DIAGNOSIS. Small bees with clear to slightly smoky wings. Males with the face entirelyyellow (sometimes the margin of the clypeus black), mark extended narrowly above theantennae; usually with marks on pronotal collar and lobes; unusual yellow mark on medianplate of forewing (Fig. 3). Lateral process of S7 longer than the width of the hemisternite,making it distinctly L-shaped; S8 dilated, without flexure, triangular. Female with longparaocular marks along eye, and a clypeal mark. Most similar to H. mimicus from Oahuand H. filicum from Hawaii; distinguishable from them by the gently arched scape and themedian plate.


Distribution. Kauai; montane mesic forest.Male. Measurements: Malar space, 3; eye, 108:100:83:58; clypeus, 53×54; frons,


Head. Malar space short (ratio about 0.2-0.3 DMO). Upper paraocular area distinctlystriate. Frons with slightly distorted polygonal pits, contiguous in center, becoming moreseparated laterally, still less than 1 pit width apart; ground minutely wrinkled and slightlylustrous in upper paraocular area, surface appearing dull between antennae (ground not

Fig. 36. Hylaeus kokeensis, new species, male (A-F) and female (G). A. Seventh sternum, ventralview. B. Eighth sternum, ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D.Genitalia, dorsal view. E. Male face with left scape. F. Male scape, lateral view. G. Female face withleft scape. 0.5 mm scale for A-D; 1 mm scale for E and G.

A B C D

0.5 mm

F

G

1.0 mm

E


evident). Lower paraocular depression very shallow, indistinct. Frontoclypeal sulcus straightor slightly arched; supraclypeal area wider than long, W/L ratios 1.1-1.5. Scape in frontalview slightly dilated, ratio about 0.6, greatest width near apex; lateral edge in frontal viewstraight; moderately arched in lateral view with maximum arch around middle; undersideof median edge with an indistinct, shallow groove; long, pale brown, simple hair alongmedian edge. Vertex hair 16-30, pale brown, mostly simple.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground slightly lustrous; hair medium-length (6-10), pale brown, weakly plumose. Scutellum with slightly larger pits, slightlylustrous. Mesepisternum with round, shallow pits about 1 pit width apart, ground smoothand lustrous, and with long, brownish white, plumose hair. Hypoepimeron with few indistinctpits, minutely granular, slightly lustrous; nearly flat. Metanotum slightly lustrous. Basalarea of propodeum with a few very short rugae at base (up to 1/3 the distance to brow,often nearly absent), the remainder including the brow smoothly coriaceous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart or more. Hair of T7 dark brown, mediumto long, erect.

Terminalia. Lateral process of S7 parallel-sided, with apex rounded, length greater thanwidth of hemisternite. Median process of S8 not or only very slightly arched, the dorsalsurface nearly in line with the sternite; strongly dilated, peaked in middle, triangular inlateral view, apex bifurcate with short, broad tips. Apices of gonoforceps subequal to penisvalves.

Color. Head and mesosoma black; metasoma dark brown. Labrum and mandible darkbrown to black. Face entirely yellow except sometimes narrow apical and lateral marginsof clypeus; paraocular marks extended narrowly above antennal sockets; supraclypeal areasometimes yellow. Pronotal lobes marked with yellow; with or without an interruptedyellow stripe on collar and marks on tegulae. Median plate of forewing yellow. Legs blackexcept for the following: front tibia with pale area and yellow mark at base; small yellowspot at apex of front femur; mid tibia narrowly yellow, and hind tibia about a third to halfyellow at base; with or without a small spot at the apex of the mid tibia anteriorly; allbasitarsi yellow, the remaining tarsi brown. Melanic individuals with basitarsal and hindtibial markings reduced to basal spots. Wings clear to slightly smoky.

Female. Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area weaklystriate above antennal sockets (much less conspicuous than in male), slightly lustrous.Frons with slightly distorted polygonal pits, contiguous in center, becoming more distantlaterally, up to 1 pit width apart near eye; ground minutely wrinkled and slightly lustrous.Middle between median ocellus and acme with a frontal line. Apex of clypeus distinctlyemarginate. Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 12-20, pale brown, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with small, shallow pits 2-3pit widths apart, ground dull; hair short (4-6), pale brown, plumose. Scutellum with slightlylarger pits, slightly lustrous. Metanotum dull. Mesepisternum with shallow pits 1-2 pitwidths apart, ground smooth, lustrous; with long, yellowish white plumose hair.Hypoepimeron weakly punctate, nearly flat. Basal area of propodeum, including brow,smoothly coriaceous and lustrous, or with a few extremely short rugae at base.


Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart or more. Hair of T6 dark brown, medium-length, erect.

Color. Clypeus with a longitudinal, apical mark occupying up to the middle third laterallyand extended back about halfway; sometimes reduced to a narrow stripe. Broad lateralmarks, not quite filling in area between clypeus and eye, extended along eye beyond antennalbases. Median plate of forewing yellow. Wings clear to slightly smoky.

Holotype ♂ labeled “Kauai, Awaawapuhi Trail, 3400', on Chamaesyce atrococca. K.Magnacca, 4 July 2000, KM-00-079” (CUIC).

Allotype ♀ with identical data (CUIC).Paratypes. Kauai, Awaawapuhi Trail, 3400', 5♀ 3-VII-2000, 4♂ 4♀ 4-VII-2000, on

Chamaesyce atrococca (KM-00-071,00-79) (CUIC); 2♂ 2♀ 4-VII-2000 (BPBM); KokeeRd., 3600', 2♀ 24-VIII-1999, on Acacia koa (KM-99-227) (CUIC).

Flower records. Acacia, Chamaesyce atrococca.Etymology. The species epithet kokeensis refers to Kokee, the region of Kauai where

the types were collected.Remarks. This species is notable for the yellow mark on the median plate at the base of

the forewing; the only other species that has this is H. kona from Hawaii, which has adistinctively shaped scape with a conspicuous groove on the underside. A large series wascollected from a single Chamaesyce bush, but it was not encountered elsewhere in thesame vicinity, except for a few individuals caught several miles away the previous year onAcacia koa. It probably occurs throughout the mesic zone in the Kokee area.

Hylaeus (Nesoprosopis) kona (Blackburn)(Fig. 37)

Prosopis kona Blackburn, 1886:144-145, 148 (♂♀; key).Nesoprosopis kona, Perkins, 1899:82 (♂♀).Hylaeus (Nesoprosopis) kona, Meade-Waldo, 1923:33 (list).Nesoprosopis kona, Daly, 1994:114 (lectotype ♂ BPBM 2595; Hawaii, western slopes

of Mauna Loa, about 6000 ft; wing length to distal end of marginal cell 3.6 mm).

DIAGNOSIS. Small bees with clear wings and with median plate at wing base yellow (Fig.3). The last is a feature shared only with H. kokeensis. Male face yellow, extended innarrow vittae above the antennal sockets; yellow marks on mandible, pronotal collar atsides and lobes, legs with tibiae and basitarsi yellow except dark brown bands on mid andhind tibiae; scape with a very distinct groove on underside, and usually a mark on thelateral edge. Female with long yellow stripe along eye, yellow marks on pronotal collar andlobes, and tegulae.

Distribution. Hawaii; montane dry forest and subalpine shrubland.Male. Measurements: Malar space, 6; eye, 105:98:91:61; clypeus, 51×51; frons,

20×22:72; interalveolar, 13:20:39:49, D 13; interocellar, 32:26:22:27, D 13; scape, 37×27;wing length, 3.8 mm (Hawaii, Kipuka Alala, 5800 ft).


Head. Malar space short to medium (ratio about 0.4-0.5 DMO). Lower half of upperparaocular area and frons striate, impunctate up to base of facial fovea; upper half withshallow pits up to 1 pit width apart near eye; frons medially with smaller, more distinct,contiguous pits and a patch of pale hair at the middle; large, lustrous impunctate areasabove the antennal sockets. Lower paraocular depression absent. Frontoclypeal sulcusstraight to slightly arched. Scape in frontal view slightly to moderately dilated, width morethan half length, ratio about 0.6-0.7; lateral edge in frontal view nearly straight, medianedge convex, reaching maximum width just past middle and narrowing slightly at apex;weakly to moderately arched in lateral view with maximum arch near middle of scape;underside of median edge with a very distinct groove; with long, yellowish white, mostlysimple hair along the median edge. Vertex hair 16-24, brownish white, weakly plumose.

0.5 mm

A B C D

E F

1.0 mm

Fig. 37. Hylaeus kona, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view, andapex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with left scape.F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shown bydashed lines in B-E.


Mesosoma. Scutum with minute pits about 2 pit widths apart, ground slightly lustrous;hair long (12-20), yellowish white to pale brown, simple. Scutellum slightly lustrous.Mesepisternum with indistinct pits, about 1 pit width apart, and long, yellowish white,plumose hair; ground minutely coriaceous, slightly lustrous. Hypoepimeron impunctate,smooth, lustrous, slightly swollen. Metanotum dull. Basal area of propodeum with ratherlong anterior area; weak, short, lineate rugae at very base; brow smoothly coriaceous andlustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous and lustrous; pale, fine,appressed hairs spaced about one hair length apart. T7 hair medium to long, brown, erect.

Terminalia. Lateral process of S7 tapered, length subequal to width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with moderately long, round tips.Apices of gonoforceps subequal to penis valves.

Color. Head and mesosoma dark brown to black; metasoma dark brown. Labrum black,mandible with a yellow spot or all yellow. Face with one large mark extended at sides toabove antennal sockets: clypeus entirely yellow or lateral margins narrowly black; paraoculararea yellow to above antennal socket in narrowed vitta; supraclypeal area to acme yellow;scape usually with a yellow spot near apex of lateral edge. Pronotum with transverse collarstripe variable, interrupted medially or with well-separated, small, lateral marks; lobes yellow.Tegula with or without a yellow spot; median plate at wing base yellow (rarely an obscureyellow-brown). Legs dark brown except: fore tibia with pale area plus long dorsal yellowstripe; mid and hind tibiae with long basal and shorter apical yellow marks, sometimesconnected by a dorsal stripe; all femora with apex yellow; all basitarsi yellow, terminaltarsal segments becoming brownish toward apex. Wings slightly smoky.

Female. Head. Malar space short (ratio about 0.4 DMO). Upper paraocular areaimpunctate and weakly striate dorsad to facial fovea, upper half with round pits 1 pit widthapart or less. Frons with round pits 1 pit width apart or less, closer in middle; very small orno impunctate area above antennae; ground minutely wrinkled, slightly lustrous; middleslightly depressed, with faint frontal line. Apex of clypeus strongly emarginate. Dorsal endof facial fovea extended to level of lateral ocellus. Vertex hair 16-26, white, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with minute, indistinct pits 1-3 pit widths apart, ground granularly coriaceous, dull; hair long (10-14), brownish white,simple. Scutellum with pits slightly larger. Mesepisternum with indistinct, shallow, nearlyround pits 1 pit width apart, and long, plumose, white hair; ground nearly smoothlycoriaceous, lustrous. Hypoepimeron impunctate, ground smoothly coriaceous, lustrous,nearly flat. Metanotum dull. Basal area of propodeum with a few short rugae at base, theremainder smoothly coriaceous, almost shining.

Metasoma. Disk of T2 impunctate, lustrous, with very short, appressed hairs spacedabout one hair length apart. T6 hair long, golden brown, erect.

Color. Head and mesosoma black; metasoma dark brown. Face with a long stripe alongthe eye, from bottom of fovea almost to base of mandible. Pronotal collar with interruptedstripe, pronotal lobe and tegula with a yellow spot; median plate yellow. Legs dark brownexcept: fore tibia with pale area plus long dorsal yellow stripe; mid and hind tibiae withlong basal and shorter apical yellow marks, sometimes connected by a dorsal stripe; all


femora with apex yellow; all basitarsi yellow, terminal tarsal segments becoming brownishtoward apex. Wings slightly smoky.

Perkins’ collections. Hawaii: Kona (specimens not located).Recent collections. Hawaii: Kipuka Alala, 5800 ft. Poikahi, 3-VI-1969, 2♀, ex Sophora

chrysophylla, C. J. Davis. Halepiula, 1♂ 3♀, 21-XI-1962, in Sophora chrysophylla, C. J.Davis.

Flower records. Bidens menziesii, Chamaesyce olowaluana, Sophora.Remarks. This species appears to be restricted to high-elevation areas of Mauna Kea

and Hualalai. It is very similar to the sympatric H. dimidiatus, with similar marks on theface, metasoma, and legs. The former has a distinctively shaped scape with the medianedge strongly convex and a conspicuous groove underneath, and marks on the mandible,median plate, and sometimes the tegula. Hylaeus dimidiatus has a nearly triangular scape,moderately dilated but with the median edge not strongly convex, and unmarked mandible,median plate, and tegula.

References. Blackburn, 1887:204-205, 208 (♂♀; key). Dalla Torre, 1896:26 (list).Perkins, 1910:603 (key). Fullaway, 1918:393 (record: Hawaii, Kilauea, Kau). Suehiro,1986:60 (record: Hawaii, Kona).

Hylaeus (Nesoprosopis) kuakea Magnacca and Daly, new species(Fig. 38)

DIAGNOSIS. Small bees with slightly smoky wings. Male with a single ivory mark on theclypeus and narrowly in the adjacent paraocular area; median frons with dense, small pits,distinctly more so than in paraocular area; hair of vertex and scutum long. Process of S8arched, very weakly dilated. Female unknown. Very similar to H. anthracinus; distinguishedby the denser, more distinct punctation of the frons, longer hairs, and generally narrowerparaocular marks.

Distribution. Oahu; lowland mesic forest.Male. Measurements: Malar space, 8; eye, 134:110:100:68; clypeus, 70×56; frons,


Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area largely impunctatealong eye, towards middle with round, shallow pits less than 1 pit width apart, extendeddown nearly to antennal sockets; ground minutely wrinkled, slightly lustrous. Frons withpits deeper, more distinct, with a large dark area of small, dense, nearly contiguous pits.Lower paraocular depression variable, distinct or not. Frontoclypeal sulcus arched. Scapein frontal view moderately dilated, W/L ratio about 0.7-0.8; lateral edge in frontal viewnearly straight in middle and rounded only at ends; weakly arched in lateral view; no grooveon underside; with medium-length, white, weakly plumose hair. Vertex hair 16-24, yellowishwhite, plumose.

Mesosoma. Scutum with distinct pits about 1 pit width apart, ground minutely granular,slightly lustrous; hair medium-length (8-12), white, plumose. Mesepisternum with distinct,


Fig. 38. Hylaeus kuakea, new species, male. A. Seventh sternum, ventral view. B. Eighth sternum,ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E.Face with left scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E.

0.5 mm

A B C D

F

1.0 mm

E

shallow, round pits, about 1 pit width apart, and long, white, weakly plumose hairs; groundminutely coriaceous, lustrous. Hypoepimeron weakly punctured, shiny, slightly swollen.Metanotum minutely wrinkled, dull. Basal area of propodeum reticulate to brow, shinybetween rugae; ground below brow minutely granular, slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, shiny; with fine, short, pale,appressed hairs spaced about one hair length apart. T7 hairs medium-length, brown,prostrate.

Terminalia. Lateral process of S7 tapered, L-shaped, length slightly greater than widthof hemisternite. Median process of S8 very weakly dilated, with a short peak at flexure,apex bifurcate with narrow tips. Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face with a single large spot:clypeus entirely ivory, lower paraocular area with a narrow stripe along clypeal suture,


supraclypeal area to acme ivory; scape unmarked. Pronotum unmarked. Legs dark brownto black, unmarked except small, obscure pale area on front tibia. Wings slightly smoky.

Female. Unknown.Holotype ♂ labeled “HI: Oahu Is, Honouliuli Preserve, Moho Gulch Ridge, 1900', over

ground. D. Hopper, 1 August 1997, KM-00-001” (CUIC).Flower records. None.Etymology. The species epithet kuakea is from the Hawaiian word meaning pale or

bleached, in reference to the white color of the facial marks.Remarks. Only two specimens are known, from a single locality in the Waianae

Mountains. One specimen is the holotype; the other was used for genetic analysis.

Hylaeus (Nesoprosopis) kukui Magnacca and Daly, new species(Fig. 39)

DIAGNOSIS. Medium-sized bees with slightly smoky wings. Male with a distinctive parallel-sided ivory stripe down the middle of the face and long hair on the scutum; moderatelydilated scape with an orange mark, unusual pale orange S6 and gonoforceps. Femaleunmarked except for prothoracic collar.

Distribution. Maui; montane wet forest.Male. Measurements: Malar space, 2; eye, 139:134:122:78; clypeus, 75×94; frons,


Head. Malar space very short (ratio about 0.1-0.2 DMO). Upper paraocular area andfrons with relatively deep, polygonal pits, almost contiguous in middle, only slightly fartherapart near eye (less than 1 pit width apart), ground minutely wrinkled, slightly lustrous; adull patch of dense and rough sculpture present about midway between each antennalsocket and lateral ocelli; middle with a fine frontal line that may be partly obscured bypunctation and sculpture. Lower paraocular depression distinct, extended above antennalsocket, shiny and striate next to eye. Frontoclypeal sulcus straight. Supraclypeal area slightlywider than long (W/L ratio 1.1-1.3). Scape in frontal view moderately dilated, ratio about0.8; lateral edge convex; moderately arched in lateral view with maximum arch near middleof scape; no groove on underside; with long, dark brown, mostly simple hair. Vertex hair20-30, dark brown, simple.

Mesosoma. Scutum with pits uniformly about 1 pit width apart, ground slightly lustrous;hair long (10-20), dark brown, mostly simple. Mesepisternum with distinct, round pitsabout 1 pit width apart, and long, pale brown, weakly plumose hair; ground smoothlycoriaceous, lustrous. Hypoepimeron weakly punctured, smooth, slightly lustrous; swollen.Metanotum dull. Basal area of propodeum strongly reticulately rugose over entire anteriorarea and brow, areolae large, ground shiny.

Metasoma. Disk of T2 with indistinct punctation, fairly densely covered with fine, dark,appressed hair, less than 1 hair length apart. Pits more distinct on T3-7. T7 hair long, darkbrown, prostrate.


Fig. 39. Hylaeus kukui, new species, male. A. Seventh sternum, ventral view. B. Eighth sternum,ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E.Face with left scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E.

C D

0.5 mm

E F

1.0 mm

A B

Terminalia. Lateral process of S7 tapered, length subequal to width of hemisternite.Median process of S8 strongly dilated, humped in lateral view, without a distinct flexure,apex bifurcate with moderately broad tips. Apices of gonoforceps subequal to penis valves.

Color. Head and mesosoma black, metasoma dark brown. Labrum and mandible blackishbrown to black. Face with a single mark: clypeus with a more or less parallel-sided ivorystripe, extended onto the supraclypeal area; mark sometimes narrow or reduced to anobscure smudge; paraocular area unmarked. Scape with an orange mark at the base of thelateral edge, absent or indistinct in some individuals with facial mark reduced. Pronotalcollar with or without an interrupted transverse ivory stripe. Legs dark brown to blackexcept: fore tibia with pale area and small yellow mark at base; fore femur orange at apex;mid tibia with yellow spot at base, with or without a small spot at the apex; hind tibia withor without an orange mark at base; all basitarsi obscure orange. S6 and ventral surface of


gonoforceps conspicuously pale orange, S6 with or without two or four obscure gray-brown spots. T7 with a large median orange spot. Wings slightly smoky.

Female. Head. Malar space very short (ratio about 0.1 DMO). Upper paraocular areaand frons with relatively deep, polygonal pits, almost contiguous in middle, only slightlyfarther apart near eye (still less than 1 pit width apart), ground minutely wrinkled, slightlylustrous; middle with a fine frontal line that may be partly obscured by punctation andsculpture. Apex of clypeus distinctly emarginate. Dorsal end of facial fovea extended tolevel of lateral ocellus. Vertex hair 16-30, dark brown, weakly plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct, evenly spacedpits about 1 pit width apart, ground dull; hair mostly medium-length (6-8, with scatteredlong hairs to 20), dark brown, simple. Mesepisternum with distinct, round pits about 1 pitwidth apart, and pale brown, mostly simple hair; ground smooth, slightly lustrous.Hypoepimeron punctured, smooth, slightly lustrous; swollen. Metanotum dull. Basal areaof propodeum reticulately rugose over entire anterior area and brow; areolae large at front,smaller around brow, ground lustrous.

Metasoma. Disk of T2 with shallow, indistinct punctation and fine, pale, appressed hair.Pits more distinct on T3-6. T6 hair long, dark brown, appressed.

Color. Body and legs dark brown to black except pronotal collar with or withoutinterrupted transverse ivory stripe, and fore tibia with obscure pale area. Wings slightlysmoky.

Holotype ♂ labeled “HI: Maui Is, Puu Kukui Trail, 3000', over ground. K. Magnacca,11 August 2000, KM-00-170” (CUIC).

Allotype ♀ with identical data (CUIC).Paratypes. Maui, Puu Kukui Trail, 3000', over ground, 3♂ 11-VIII-2000 (KM-00-170)

(CUIC). Kipahulu Valley, Camp 2, 1250 m, 7♂ 1♀ 13-17-VIII-1967, N. Wilson; Hana,10♂ 2♀ 7-V-1920, E. H. Bryan Jr. (BPBM).

Flower records. None.Etymology. The species epithet kukui refers to the summit of West Maui, named after

the candlenut tree.Remarks. This species is related to H. muranus of Hawaii, but lacks the transverse

scape and any trace of iridescence on the abdomen. All of KM’s specimens were caughtflying in a small area under a low fern; a long series from East Maui exists at the BishopMuseum. It appears to occur widely in the wet forests of Maui.

Hylaeus (Nesoprosopis) laetus (Perkins)(Fig. 40)

Nesoprosopis laeta Perkins, 1899:81-82 (♂♀).Hylaeus (Nesoprosopis) laetus, Meade-Waldo, 1923:33 (list).Nesoprosopis laeta, Daly, 1994:117 (lectotype ♂ BMNH 17a 134a; Oahu, Waianae

Mts.; wing length, 4.0 mm).

DIAGNOSIS. Medium-sized bees with clear to dark smoky wings. Male face usually withoval yellow mark extending onto paraocular area, or mark on clypeus only, scape slightly


dilated and moderately to strongly arched, legs marked with yellow, lateral process of S7longer than width of hemisternite; median process of S8 not dilated and with long, narrowtips, apices of gonoforceps subequal to penis valves. Female black with pronotal lobesyellow or unmarked; hair of scutum short. Sister species of H. difficilis; see remarks forthat species.

Distribution. Hawaii, Maui, Lanai, Oahu, and Kauai; found primarily in both lowlandand montane dry shrubland, less commonly in mesic and wet forest.

Male. Measurements: Malar space, 5; eye, 124:110:100:72; clypeus, 65×64; frons,19×24:83; interalveolar, 15:23:50:59, D 19; interocellar, 32:28:22:28, D 15; scape, 39×20;wing length, 4.3 mm (Oahu, Waianae Mts., 2000 ft, BPBM 3471).

0.5 mm

A B C D

E F

1.0 mm

Fig. 40. Hylaeus laetus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.


Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area with indistinct,round, shallow pits 1 pit width apart or less (some specimens with pits almost totallyobscured by minutely wrinkled, slightly lustrous ground; other specimens with relativelysmooth lustrous ground); dense, minute, parallel wrinkles below punctured area and alongthe eye, and ground rather smooth and lustrous; frons with pits closer, more distinct,contiguous medially, slightly distorted; fine frontal line present in some specimens; shiny,median, depressed area just above prominent acme. Lower paraocular depression distinct,but shallow. Frontoclypeal sulcus slightly arched. Scape in frontal view slightly dilated,width about half length, ratio about 0.5; lateral edge in frontal view nearly straight orslightly concave; moderately to strongly arched in lateral view with maximum arch nearmiddle of scape; width of scape in lateral view more or less uniform in basal half, expandedgradually in width in apical half; no groove on underside; with medium-length, pale brown,weakly plumose hair. Vertex hair 12-20, pale brown to brown, plumose.

Mesosoma. Scutum with small, shallow pits about 2 pit widths apart, ground slightlylustrous; hair medium-length (8-12), white to pale brown, plumose. Scutellum with pitsmore distinct, ground lustrous. Mesepisternum with indistinct to barely discernible pits,about 1-2 pit widths apart, and long, brownish to yellowish white, plumose hair; groundminutely coriaceous, slightly lustrous. Hypoepimeron impunctate, lustrous, nearly flat.Metanotum dull. Basal area of propodeum lineately rugose about halfway or more to brow,spaced 2-4 ruga widths apart or more, often shiny between rugae; brow granular to relativelysmoothly coriaceous, dull to slightly lustrous.

Metasoma. Disk of T2 impunctate; smoothly coriaceous, lustrous, with pale, short,appressed hairs spaced more than a hair length apart. T7 hair short, yellowish white to red-brown, prostrate.

Terminalia. Lateral process of S7 tapered, apex translucent, length greater than widthof hemisternite. Median process of S8 linear and with a slight triangular dilation at tip, andrather long, very slender, cuticular bifurcations with hairs (the slender bifurcations easilyconfused with hairs at tip of process or may be broken and lost). Apices of gonoforcepssubequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face with a single large spot:clypeus entirely yellow or, if other marks missing, then with only a central spot; lowerparaocular area yellow along sulci below antennal socket, or unmarked; supraclypeal areato acme yellow or unmarked; scape unmarked. Pronotum with lobes yellow, or only anapical dot, or unmarked. Legs dark brown to black except pale area on fore tibia andvariable amounts of the following yellow marks: fore leg with basal stripe up to most oflength of tibia; basitarsus mostly yellow or at least a basal trace; and terminal tarsal segmentsyellow to brown. Mid leg with basal stripe up to half length of tibia and sometimes combinedwith a subapical mark; basitarsus mostly yellow or at least a basal trace; and terminal tarsalsegments yellow to brown. Hind leg with tibia entirely yellow in basal half, sometimescombined with subapical mark, or yellow extended full length in frontal view and in caudalview, dark in apical half; basitarsus mostly yellow or at least a basal trace; and terminaltarsal segments yellow to brown. Wings clear to smoky.


Female. Head. Malar space short to medium (ratio about 0.3 to 0.5 DMO). Upperparaocular area with indistinct, round, shallow pits 1 pit width apart or less; ground minutelywrinkled, slightly lustrous; in some specimens, frons with pits closer, medially contiguous,slightly distorted; fine frontal line sometimes present. Apex of clypeus shallowly emarginate.Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 6-14(-20), brown,plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with shallow pits about 2 pitwidths apart, ground slightly lustrous; hair medium-length (6-10), yellowish white, plumose.Mesepisternum with indistinct to barely discernible pits, about 2 pit widths apart, andmedium-length, yellowish white, plumose hair; ground minutely coriaceous or slightly striate,lustrous. Hypoepimeron impunctate, lustrous, nearly flat. Metanotum slightly lustrous.Basal area of propodeum coarsely lineately rugose about halfway to brow, spaced 2-4 rugawidths apart, rugae weaker near brow; brow smoothly coriaceous, lustrous.

Metasoma. Disk of T2 impunctate; smoothly coriaceous, lustrous, with pale, short,appressed hairs spaced more than a hair length apart. T6 hair medium-length, light brown,prostrate.

Color. Head and mesosoma black; metasoma dark brown. Face unmarked. Pronotumwith lobes yellow or unmarked. Legs dark brown to black, unmarked or with obscure ordistinct basal yellow spots on tibiae, largest on hind tibia. Wings smoky to dark smoky.

Perkins’ collections. Hawaii: Kilauea (BMNH). Lanai: Koele, 2000 ft (OUMNH);“Lanai” (BPBM). Molokai: “Molokai” (BPBM); Molokai Mts., 3000 ft (BMNH, BPBM);Plains (BMNH, BPBM). Oahu: Honolulu, 2000 ft (BPBM); Honolulu Mts., 1500 ft(BPBM); Kaala, 2000 ft (OUMNH); Tantalus (BPBM); Waianae (BMNH), coast (BPBM),coast, field note #672 (OUMNH); Waianae Mts. (BPBM), 2000 ft (BMNH, BPBM,OUMNH). Kauai: Halemanu, 4000 ft (field note #501) (BMNH, BPBM, OUMNH);“Kauai” (OUMNH); Makaweli, >2000 ft (BPBM), 2000 ft (BMNH, BPBM); Waimea,2000 ft (BMNH, BPBM), >2000 ft (BPBM).

Recent collections. Hawaii: Hilina Pali Rd., 2000 ft; Kipuka Nene, 2900 ft; Puu Waawaa,4100 ft; Kipuka Alala, 5800 ft; Humuula Saddle, 6800 ft. Maui: Lahainaluna, 1800 ft;Kahoma, 2100 ft. Lanai: Kahue, 1400 ft; Garden of the Gods, 1400 ft. Oahu: PaholeNatural Area Reserve, 2000 ft. Kauai: Alakai Swamp Trail, 3600 and 4000 ft; Kokee Rd.,3600 ft; Nualolo Cliff Trail, 2800 ft; Paaiki Valley, 2700 ft.

Flower records. Acacia, Bidens menziesii, B. sandvicensis, Chamaesyce atrococca, C.multiformis, C. olowaluana, Claoxylon, Dodonaea, Metrosideros, Myoporum, Plantagolanceolata*, Santalum freycinetianum, S. paniculatum, Scaevola procera, Sesbaniatomentosa, Sida, Sophora, Styphelia.

Remarks. This species occurs in many of the same habitats as H. difficilis, but is morecommonly found at lower elevations (below 4000 ft). See remarks for H. difficilis.

References. Perkins, 1899:105 (host of cleptoparasite: N. hostilis); 1910:602 (key);1913:82 (biogeography). Fullaway, 1918:393 (record: Hawaii, Kau, Kilauea). Illingworth,1928:46 (record: Lanai).


Hylaeus (Nesoprosopis) longiceps (Perkins)(Fig. 41)

Nesoprosopis longiceps Perkins, 1899:98 (♂♀; Pl. 2, Fig. 3).Hylaeus (Nesoprosopis) longiceps, Meade-Waldo, 1923:34 (list).Nesoprosopis longiceps, Daly, 1994:118 (lectotype ♂ BMNH 17a 160ab; Maui, Wailuku;

wing length, 4.3 mm).

DIAGNOSIS. Medium-sized bees with clear to slightly smoky wings. Male lower faceentirely yellow, extended at sides in a broad stripe above antennal sockets, unusualsupraclypeal area very long and narrow, scape weakly dilated (H. solaris from Kauai has asimilar long supraclypeal area, but the scape is dilated). Female black and unmarked.

Distribution. Maui, Molokai, Lanai, and Oahu; coast and dry lowlands.Male. Measurements: Malar space, 6; eye, 159:115:102:74; clypeus, 79×69; frons,

31×22:105; interalveolar, 13:23:58:67, D 23; interocellar, 35:26:20:28, D 18; scape, 51×20;wing length, 4.2 mm (Maui, Wailuku, BPBM 3508).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area with distinct,round pits 1-2 pit widths apart, ground nearly smooth, slightly lustrous; frons with pits stilldistinct, smaller, contiguous and distorted, and merging at the middle into a localizeddepression of dull, rough ground above the antennal sockets; otherwise no special sculpture,frontal line not evident; impunctate area above antennal sockets very small. Lower paraoculardepression very shallow or absent. Frontoclypeal sulcus straight. Scape in frontal view notdilated or very weakly dilated, width less than half length, ratio about 0.3-0.4; lateral edgein frontal view nearly straight or slightly concave; slightly arched in lateral view withmaximum arch near middle of scape; no groove on underside; with short, yellowish white,plumose hair. Vertex hair 8-16, yellowish white, plumose.

Mesosoma. Scutum with distinct pits about 1 pit width apart, ground nearly smooth,slightly lustrous; hair medium-length (8-10), white to pale brown, plumose. Scutellumwith pits slightly larger in some bees, lustrous. Mesepisternum with shallow pits about 1pit width apart, and medium-length, white, plumose hair; ground nearly smooth, lustrous.Hypoepimeron large, impunctate or obscurely punctured, smooth, lustrous; slightly swollen.Metanotum dull to slightly lustrous. Basal area of propodeum lineately rugose to brow,shiny between rugae, brow nearly smooth, lustrous; varying to reticulately rugose to brow,large shiny areolae at front, brow weaker reticulately rugose, shiny between rugae.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with very short, fine,pale, appressed hairs spaced about one hair length apart or less. T7 hair short, goldenbrown, appressed.

Terminalia. Lateral process of S7 tapered, pale, length greater than width of hemisternite.Median process of S8 weakly dilated and rather thick, apex bifurcate with long, narrow,rounded, sparsely hairy tips. Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black, labrum with or without a yellowspot. Face with one large mark: clypeus entirely yellow or nearly so; paraocular area yellowto above antennal socket; supraclypeal area to acme yellow; scape unmarked or with ayellow spot or line on the lateral edge. Pronotum unmarked or with a yellow spot on lobe;


tegula with or without a yellow spot. Legs dark brown to blackish brown except: fore tibiawith pale area and yellow spots at base and apex that may be connected by a dorsal yellowishstripe, tarsus yellow, brownish apically; mid and hind tibiae with small yellow marks atbase and apex; mid and hind tarsi yellow, brownish apically. Wings clear.

Female. Head. Malar space medium (ratio about 0.6 DMO). Upper paraocular areawith distinct, round pits 1-2 pit widths apart, ground nearly smooth, slightly lustrous; fronswith pits still distinct, medially smaller, contiguous, and distorted; at the middle, with orwithout a few longitudinal wrinkles or a faint, fine frontal line. Apex of clypeus shallowlyemarginate. Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 6-8,pale brown, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits about 1 pitwidth apart, ground nearly smooth, slightly lustrous; hair short (4, relatively even, occasional

0.5 mm

A B C D

E F

1.0 mm

Fig. 41. Hylaeus longiceps, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


hairs to 8), white, plumose. Scutellum with pits slightly larger in some bees, lustrous.Mesepisternum with shallow pits about 1 pit width apart, and yellowish white, weaklyplumose hair; ground nearly smooth, lustrous. Hypoepimeron large, impunctate, smoothto very weakly striate, lustrous; slightly swollen. Metanotum dull to slightly lustrous. Basalarea of propodeum lineately rugose or reticulate to brow, shiny between rugae, some beeswith sparse rugae; brow nearly smooth to granularly coriaceous, slightly lustrous to dull.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with short or veryshort, fine, pale, appressed hairs spaced a hair length apart or more. T6 hair long, goldenbrown, appressed.

Color. Head and mesosoma brown to black; metasoma brown to dark blackish brown.Face and pronotum unmarked. Legs dark brown; unmarked or with a small yellow spot atbase of front tibia. Wings nearly clear to slightly smoky.

Perkins’ collections. Maui: Haleakala, 2000 ft (BPBM); “Maui” (BPBM); Wailuku(BMNH, BPBM, OUMNH); sand hills (BPBM). Lanai: “Lanai” (BPBM); Manele (BMNH,BPBM). Molokai: Kaunakakai (OUMNH); Molokai coast and plains (BMNH, BPBM);Molokai Mts. (BPBM); West end of Molokai (BPBM). Oahu: Waianae (OUMNH).

Recent collections. Maui: Waiehu dune, 200 ft. Lanai: Kahue, 1400 ft; Polihua Rd.,1000 ft; Shipwreck Beach, coast. Molokai: Moomomi, coast. Oahu: Kaena Point NaturalArea Reserve, coast.

Flower records. Chamaesyce degeneri, Myoporum, Santalum ellipticum, Scaevolacoriacea (Maui), S. sericea, Sesbania tomentosa, Sida, Vitex rotundifolia.

Remarks. The supraclypeal area is variable in shape, but consistently longer than wide(8 bees, W/L ratios 0.6-0.8), and the face is unusually long and narrow overall. Only H.solaris has a similarly narrow face; while it also has similar facial markings, the two speciesare not closely related.

References. Perkins, 1910:602 (key); 1911:727 (Blackburn had series of mixed species);1913:79 (morphology). Fullaway, 1918:396 (record: Oahu, Makapuu; Lanai).

Hylaeus (Nesoprosopis) mana Magnacca and Daly, new species(Fig. 42)

DIAGNOSIS. Very small bees with clear wings. Male face nearly all yellow, only supraclypealarea and narrow margins of clypeus black, the yellow marks extended above antennalsockets; scape weakly dilated, about twice as long as wide, weakly arched. Median processof S8 swollen at base, the remainder weakly dilated and strongly arched. Female with long,broad paraocular marks, a transverse clypeal mark, and marks on pronotal collar, lobes,and tegulae. Both sexes with extensive yellow marks on legs.

Distribution. Oahu; mesic forest. Known from only one collection.Male. Measurements: Malar space, 2; eye, 80:76:68:46; clypeus, 38×42; frons, 12×14:60;

interalveolar, 12:16:36:42, D 12; interocellar, 22:20:12:22, D 10; scape, 30×14; wing length,2.9 mm (holotype).

Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular area with distinctpits about 1 pit width apart, denser and almost touching in the median frons; ground strongly


lustrous in upper paraocular area, less so on frons; a large dark patch between acme andmedian ocellus formed by dense punctation and dull sculpture, covered with pale, appressedhairs. Lower paraocular depression very shallow, indistinct. Frontoclypeal sulcus slightlyarched or nearly straight; supraclypeal area wider than long, W/L ratio 1.2-1.4. Scape infrontal view hardly dilated, ratio about 0.5, greatest width near apex; lateral edge in frontalview mostly straight, with a slight knob at the apex; weakly arched in lateral view; undersideof median edge with an indistinct, shallow groove; long, pale, simple hair along medianedge. Vertex hair 8-12, pale brown, simple.

Mesosoma. Scutum with shallow pits 2-3 pit widths apart, ground minutely granular,slightly lustrous; hair uneven, short to medium-length (4-12), pale, mostly simple. Scutellumsmoother, more lustrous. Mesepisternum with shallow, indistinct pits about 1-3 pit widthsapart, ground strongly lustrous, with long, white, plumose hair. Hypoepimeron impunctate,shiny, slightly swollen. Metanotum slightly lustrous. Basal area of propodeum with a fewvery short rugae at base or none, the remainder including the brow smoothly coriaceous,lustrous; dorsal surface long, as long as scutellum. Lateral carinae indistinct, evident onlyposteriorly.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,prostrate hairs spaced more than a hair length apart. Hair of T7 brown, medium to long,erect.

Terminalia. Lateral process of S7 parallel-sided, length greater than width of hemisternite;hemisternite L-shaped. Median process of S8 swollen at base, the remainder narrow andstrongly arched, apex at about the level of the disk of S8 and bifurcate, with long, strapliketips. Apices of gonoforceps exceeding penis valves, inwardly pointed.

Color. Labrum and mandible dark brown to black. Face with three nearly contiguousspots: clypeus nearly all yellow except narrow margins of clypeus; paraocular marks extendednarrowly above antennal sockets; supraclypeal area black. Pronotum with an interruptedtransverse stripe on collar and marks on lobes and tegulae. Legs black except: front tibiawith large pale area and a dorsal yellow stripe the entire length; mid tibia with basal quarterto third, and apical quarter, yellow; hind tibia with basal half and narrow apical ring yellow;marks on apices of all femora; all basitarsi yellow. Wings clear.

Female. Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area andfrons with evenly spaced pits, about 1 pit width apart even at center below median ocellus,but with a small area of slightly denser punctation halfway between acme and ocellus;ground almost smooth, strongly lustrous. Middle between median ocellus and acme with afaint frontal line. Apex of clypeus distinctly emarginate. Dorsal end of facial fovea notreaching level of lateral ocellus. Vertex hair 6-8, pale brown, simple.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with shallow pits 2-3 pitwidths apart, ground minutely granular and weakly lustrous; hair short (2-6; but specimenabraded), pale, simple. Scutellum smoother, more lustrous. Metanotum dull. Mesepisternumwith shallow, indistinct pits 1-2 pit widths apart, ground smooth, strongly lustrous; withmedium-length, white, weakly plumose hair. Hypoepimeron impunctate, strongly lustrous,slightly swollen. Basal area of propodeum with a few very short rugae at base or none, theremainder including the brow smoothly coriaceous, lustrous; dorsal surface long, as longas scutellum. Lateral carinae indistinct, evident only posteriorly.


Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,prostrate hairs spaced about one hair length apart or more. Hair of T6 long, brown, erect.

Color. Head and mesosoma black; metasoma dark brown. Face marked with yellow:clypeus with a transverse apical mark (possibly sometimes absent); paraocular area with abroad stripe, extended along eye up to antennal socket. Pronotum with marks on lobes andinterrupted transverse collar stripe; tegula with a yellow spot. Legs black except: fronttibia with large pale area and a dorsal yellow stripe half the length; mid tibia with basal andapical quarters yellow; hind tibia with basal half and narrow apical ring yellow; marks on

Fig. 42. Hylaeus mana, new species, male (A-F) and female (G). A. Seventh sternum, ventral view.B. Eighth sternum, ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia,dorsal view. E. Male face with left scape. F. Male scape, lateral view. G. Female face with left scape.0.5 mm scale for A-D; 1 mm scale for E and G.

A B C D

0.5 mm

1.0 mm

G

F

E


apices of all femora; all of front and base of middle basitarsus yellow. Wings clear. Metasomalterga and sterna 1-5 with thin cuticle apically and laterally, creating the appearance oforange bands.

Holotype ♂ labeled “Oahu, Manana Trail, 1400', on Santalum freycinetianum. K.Magnacca, 3 Mar 2002, KM-02-043” (CUIC).

Allotype ♀ with identical data (CUIC).Paratype. 1♂ with identical data (BPBM).Flower records. Santalum freycinetianum.Etymology. The species epithet mana is from the Hawaiian word meaning forked,

referring to the facial marks; it also means spiritual power, which this tiny creature mustpossess to have made itself known through a chance collection shortly before publicationof this volume.

Remarks. This is the smallest species of Hawaiian Hylaeus. The only collection camefrom a habitat (mesic koa forest) and plant (Santalum) where few bees have been found onOahu. The weakly arched scape with a poorly developed groove suggests that it may berelated to H. dumetorum of Hawaii. Males can immediately be distinguished from thesympatric H. mimicus and H. specularis by the shape of S8, which is hardly dilated andstrongly arched; all three males of H. mana have similar facial marks with more yellowthan either of those species. Females can be distinguished by the evenly, more widely spacedpits of the frons, and the transverse rather than longitudinal clypeal mark (H. specularismay occasionally have a transverse clypeal mark as well).

Hylaeus (Nesoprosopis) mauiensis (Perkins)(Fig. 43)

Nesoprosopis mauiensis Perkins, 1899:94-95 (♂♀).Hylaeus (Nesoprosopis) mauiensis, Meade-Waldo, 1923:34 (list).Nesoprosopis mauiensis, Daly, 1994:118 (lectotype ♂ BMNH 17a 148; Maui, Haleakala,

5000 ft).

DIAGNOSIS. Small bees with clear wings. Male with small, irregular yellow marks only inparaocular areas; legs unmarked; basal area of propodeum with rather long base and atvery front with short, linear rugae spaced about 3 rugae apart; brow smooth. Femaledescribed by Perkins (1899:95) had a narrow yellow line along eye margins, yellow marksat each side of collar, and basal area of propodeum unsculptured.

Distribution. Maui; montane wet forest. No recent collections.Male. Measurements: Malar space, 3; eye, 92:92:83:54; clypeus, 45×40; frons, 15×18:69;

interalveolar, 14:18:40:43, D 17; interocellar, 26:28:18:28, D 15; scape, 32×23; wing length,3.6 mm (H. mauiensis lectotype).

Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular area with shallow,round pits about 1 pit width apart, ground minutely roughened with parallel wrinkles curvedfrom near median ocellus to along eye; on frons pits closer and contiguous; at middle pitsdense, contiguous, indistinct, distorted, and with a small dark spot of hairs just above and


between impunctate areas above the antennal sockets; no median groove. Lower paraoculardepression absent. Frontoclypeal sulcus straight. Scape moderately dilated, width morethan half length, ratio about 0.7; lateral edge in frontal view straight; moderately arched inlateral view with maximum arch near middle of scape.

Mesosoma. Scutum with indistinct, minute pits, 2-3 pit widths apart, ground granular,slightly lustrous. Scutellum smooth, lustrous. Mesepisternum largely impunctate with sparse,simple hairs less than a hair length apart, ground smooth, lustrous. Hypoepimeronimpunctate, smooth, lustrous; slightly swollen. Metanotum mostly smooth, slightly lustrous.Basal area of propodeum with rather long base; at very front with short, linear rugaespaced about 3 rugae apart; brow smoothly coriaceous and lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; mostly bare, otherwisewith fine, pale, appressed hairs spaced about one hair length apart.

Terminalia. Lateral process of S7 parallel-sided, with length greater than width ofhemisternite. Median process of S8 moderately dilated, apex bifurcate with broad, roundedtips of moderate length. Apices of gonoforceps subequal to penis valves.

Color. Head and mesosoma black; metasoma dark brown. Face with irregular yellowstripes in paraocular areas not reaching level of antennal sockets; clypeus, frons and scapeunmarked. Pronotum unmarked. Legs dark brown except fore tibia with pale area. Wingsclear.

Female. Specimen not located. Described by Perkins (1899:95) as follows. “Female,with a narrow yellow line along the margins of the eyes, and one on each side of the hindmargin of the prothorax yellow; head above the antennae with the surface shining (exceptnarrowly in the middle), finely and closely punctured; mesothorax dull, and very finelypunctured; scutellum also dull, finely and somewhat remotely punctured; anterior area of

Fig. 43. Hylaeus mauiensis, male. A. Face with left scape. B. Scape, lateral view.

A B


the propodeum with only the fine general surface rugulosity and shining in certain aspects;wings clear and iridescent.”

Perkins’ collections. Maui: Haleakala, 5000 ft (BMNH).Recent collections. None.Flower records. None.Remarks. Known only from the male type specimen and the description of a female.References. Perkins, 1910:603 (key).

Hylaeus (Nesoprosopis) melanothrix (Perkins)(Fig. 44)

Nesoprosopis melanothrix Perkins, 1899:86-87 (♂♀; Pl. 2, Fig. 14; nest in Metrosiderosand Psychotria [referred to as Straussia]).

Hylaeus (Nesoprosopis) melanothrix, Meade-Waldo, 1923:34 (list).Nesoprosopis melanothrix, Daly, 1994:118 (lectotype ♂ BMNH 17a 119b; Maui,

Haleakala, 5000 ft; wing length, 5.4 mm).

DIAGNOSIS. Medium-sized bees with dark smoky wings; long brown body hair; unusualpropodeum with basal area strongly sloping, brow indistinct, and sculpture, if any, weak atvery front, otherwise smooth, lustrous. Male with unique scape deeply arched with lateralapex expanded. Female black and unmarked; long malar space in comparison to relativelysmall ocellus (0.8 DMO).

Distribution. Maui; montane wet forest. No recent collections.Male. Measurements: Malar space, 6; eye, 120:123:110:70; clypeus, 61×59; frons,

23×26:87; interalveolar, 18:26:50:54, D 20; interocellar, 33:36:20:36, D 14; scape, 44×30;wing length, 4.5 mm (Maui, Haleakala, 5000 ft, BPBM 3509).

Head. Malar space short (ratio about 0.4 DMO). Apex of clypeus shallowly emarginate.Upper paraocular area and frons with pits less than 1 pit width apart, crowded and distortedin subpolygonal shapes, and at the middle a patch of long, dark hairs, but no dark rhomboidspot of dense hairs; shiny area between patch and antennal sockets. Lower paraoculardepression absent. Frontoclypeal sulcus slightly arched. Scape in frontal view moderatelydilated, width more than half length, ratio about 0.7; lateral edge in frontal view slightly todistinctly concave; strongly arched in lateral view with maximum arch near apex of scape;underside of median edge with a groove; with medium-length, brown, simple hair on thefrontal surface. Vertex hair 20-40, dark brown, simple.

Mesosoma. Scutum with shallow pits 1-2 pit widths apart (pits less distinct on somebees), ground slightly lustrous; hair long (14-28), simple, brown. Scutellum with pits slightlylarger, ground slightly lustrous. Mesepisternum with indistinct pits, about 1 pit width apart,and long, brownish, weakly plumose hair; ground minutely coriaceous, lustrous.Hypoepimeron impunctate, lustrous, slightly swollen. Metanotum dull. Basal area ofpropodeum strongly sloping, nearly glabrous or with sparse, weak, lineate rugae at base;brow indistinct, smoothly coriaceous and lustrous.


Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart. T7 hair medium-length, brown, erect.

Terminalia. Lateral process of S7 tapered, with length less than width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with rounded tips. Apices ofgonoforceps subequal to penis valves.

Color. Head and mesosoma dark brown to black; metasoma dark brown. Labrum andmandible blackish brown to black. Face with three separate marks: clypeus with irregular,subapical yellow spot or at least a trace; lower paraocular area with irregular yellow stripealong the eye below antennal socket; supraclypeal area and scape unmarked. Pronotumunmarked. Legs dark brown to black; unmarked except pale area on fore tibia. Wings darksmoky.

0.5 mm

A B DC

E F

1.0 mm

Fig. 44. Hylaeus melanothrix, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


Female. Head. Malar space long in comparison to relatively small ocellus (ratio about0.8 DMO). Upper paraocular area and frons convex; upper paraocular area with nearlyround pits less than 1 pit width apart, ground lustrous to shiny; frons with pits crowdedmedially and partly distorted in shape, and at the middle a few, fine, longitudinal carinulae;frontal line distinct above acme; small shiny area just above antennal socket. Apex of clypeusshallowly to deeply emarginate. Dorsal end of facial fovea extended to level of lateralocellus. Vertex hair 20-40, dark brown, weakly plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground lustrous; hair medium-length (8-16), mostly simple, brown. Scutellum similar,pits slightly larger. Mesepisternum with distinct pits, about 1-2 pit widths apart, and long,brownish white, weakly plumose hair; ground minutely coriaceous, lustrous. Hypoepimeronimpunctate, lustrous, nearly flat. Metanotum dull. Basal area of propodeum strongly sloping,nearly glabrous or with sparse, weak, lineate rugae at base; brow indistinct, smoothlycoriaceous and lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, but polished and shiny, withsparse, fine, pale, appressed hairs spaced more than a hair length apart. T6 hair long, darkbrown, erect.

Color. Body and legs black; unmarked. Wings dark brown.Perkins’ collections. Maui: Haleakala, 4000 ft (BMNH), 5000 ft (BMNH, BPBM,

OUMNH), >5000 ft (BPBM).Recent collections. None.Flower records. None.Remarks. The measured male has the extreme ratio “interocellar distance/diameter of

median ocellus” at 2.4, indicating that the lateral ocelli are far apart relative to the diameterof the median ocellus. However, the ratio “ocellus-eye/interocellar distance” at 1.1, indicatingthe position of the lateral ocelli relative to the eye, is not at all exceptional.

References. Perkins, 1910:603 (key); 1913:79 (nest).

Hylaeus (Nesoprosopis) mimicus Magnacca and Daly, new species(Fig. 45)

DIAGNOSIS. Small bees with slightly smoky wings. Male face with three marks separatedby dark margins of clypeus, extended narrowly along eye; scape strongly arched, almostangled, with a groove underneath. Female with long, broad paraocular marks; a triangularclypeal mark; and marks on pronotal collar, lobes, and tegulae. See remarks for similarspecies.

Distribution. Oahu; montane mesic and wet forest.Male. Measurements: Malar space, 3; eye, 102:96:83:57; clypeus, 54×47; frons,


Head. Malar space short (ratio about 0.2 DMO). Upper paraocular area and frons withslightly distorted pits, very small and nearly contiguous in center; ground minutely wrinkled


and lustrous in upper paraocular area, less so on frons; large dark patch between antennaeand median ocellus formed by dense punctation and sculpture. Lower paraocular depressionvery shallow, indistinct. Frontoclypeal sulcus arched; supraclypeal area much wider thanlong, W/L ratio 1.2-1.6. Scape in frontal view slightly dilated, ratio about 0.6, greatestwidth near apex; lateral edge in frontal view slightly concave; strongly arched in lateralview with maximum arch around middle; underside of median edge with a groove; medium-length, pale brown, simple hair along median edge. Vertex hair 16-24, brown, simple.

Mesosoma. Scutum with shallow, indistinct pits 2-3 pit widths apart, ground granular,dull; hair medium-length (8-12), pale brown, mostly simple. Scutellum with slightly larger

Fig. 45. Hylaeus mimicus, new species, male (A-F) and female (G). A. Seventh sternum, ventralview. B. Eighth sternum, ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D.Genitalia, dorsal view. E. Male face with left scape. F. Male scape, lateral view. G. Female face withleft scape. 0.5 mm scale for A-D; 1 mm scale for E and G.

A B C D

0.5 mm

E

F

G

1.0 mm


pits, slightly lustrous. Mesepisternum with round, shallow pits about 1-2 pit widths apart,ground smooth and lustrous, with long, brownish white, plumose hair. Metanotum slightlylustrous. Basal area of propodeum with few very short rugae at base or none, the remainderincluding the brow smoothly coriaceous, lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced more than a hair length apart. Hair of T7 black, medium-length,erect.

Terminalia. Lateral process of S7 parallel-sided, with length greater than width ofhemisternite. Median process of S8 moderately dilated, with a distinct flexure, apex bifurcatewith rounded tips. Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible dark brown to black. Face with three separate spots: clypeuslargely yellow except broad basal and lateral margins; paraocular marks extended narrowlyabove antennal sockets; supraclypeal area black. Pronotum variable, with or withoutinterrupted transverse stripe on collar and marks on lobes. Legs black except: front tibiawith large pale area; all tibiae with yellow spots at bases of tibiae; base of fore basitarsusand apex of femur with yellow marks; mid and hind basitarsi with or without marks ofvarious sizes. Wings slightly smoky.

Female. Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area andfrons with slightly distorted polygonal pits, nearly contiguous in center, more separatedlaterally, up to 1 pit width apart near eye; ground minutely wrinkled, somewhat obscuringpits; slightly lustrous. Middle between median ocellus and acme with an obscure frontalline. Apex of clypeus distinctly emarginate. Dorsal end of facial fovea extended to level oflateral ocellus. Vertex hair 16-24, brown, mixed simple and plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with small, shallow pits 2-3pit widths apart, ground dull; hair medium (8-10), pale brown, simple. Scutellum withslightly larger pits, slightly lustrous. Metanotum dull. Mesepisternum with shallow pits 1-2 pit widths apart, ground smoothly coriaceous, lustrous; with long, yellowish white, plumosehair. Hypoepimeron weakly punctate, nearly flat. Basal area of propodeum, including brow,smoothly coriaceous and lustrous (brow almost shining), with or without a few extremelyshort rugae at base.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart or more. T6 hair long, dark brown,erect.

Color. Head and mesosoma black; metasoma dark brown. Face with three separate yellowmarks: clypeus with a more or less triangular apical mark, occupying about the middle halflaterally and extended back about halfway; paraocular area with a broad stripe, not quitefilling in area between clypeus and eye, extended along eye beyond antennal socket.Pronotum with marks on lobes and interrupted transverse collar stripe; tegula with a yellowspot. Legs black except: front tibia with pale area and dorsal yellow stripe, front basitarsusand apex of front femur yellow; mid and hind tibiae with yellow spots at base; mid and hindbasitarsi with or without yellow marks. Wings slightly smoky.

Holotype ♂ labeled “Oahu, Aiea Trail, 1700', on Acacia koa. K. Magnacca, 2 Jun 2000,KM-00-012” (CUIC).


Allotype ♀ labeled “Oahu, Aiea Trail, 1900', on Psychotria mariniana. K. Magnacca,14 Aug 2000, KM-00-179” (CUIC).

Paratypes. Oahu, Aiea Trail, 1♂ 14-VIII-2000, on Ilex anomala (KM-00-175); 4♂ 14-VIII-2000, on Psychotria mariniana (KM-00-179) (CUIC). Aiea Trail, 3♂ 14-VIII-2000,on Scaevola gaudichaudiana (KM-00-173); Wiliwilinui Trail, 1800', 2♂ 1♀ 26-VII-1999,on Scaevola gaudichaudiana (KM-99-167); Opaeula, 1♂ 30-III-1913, O. H. Swezey(BPBM).

Flower records. Acacia, Ilex anomala, Metrosideros, Psychotria mariniana, Scaevolagaudichaudiana.

Etymology. The species epithet mimicus refers to the close resemblance of this speciesto H. filicum on distant Hawaii.

Remarks. This species may be responsible for purported records of H. filicum fromOahu, as it is very similar. It can be distinguished by the much shorter hair on the mesonotumand vertex (that of H. filicum measures 12-20). It also lacks the unusual bulge at the apexof the scape that causes that of H. filicum to be much more strongly arched on the lowersurface than the upper. The upper frons of H. mimicus appears dark due to close sculpture,but it lacks the conspicuous rhomboid patch of dark hairs found in H. filicum and otherrelated species. The amount of yellow on the face of the female is unusual, and similar tothat of H. kokeensis, which has a yellow median plate in both sexes. It is widely present onleeward ridges of the Koolau Mountains, but appears to be less abundant than H. connectensand H. unicus, the two species most commonly collected there.

Hylaeus (Nesoprosopis) muranus (Warncke), new combination(Fig. 46)

Nesoprosopis insignis Perkins, 1899:110-111 (♂♀; Pl. 2, Fig. 24).Hylaeus (Nesoprosopis) insignis, Meade-Waldo, 1923:35 (list).Prosopis murana Warncke, 1970:747 (replacement name for Nesoprosopis insignis

Perkins, junior homonym of Hylaeus insignis Forster)Nesoprosopis insignis, Daly, 1994:117 (lectotype ♂ BMNH 17a 144b; Hawaii, Kau,

4000 ft; wing length, 6.3 mm, tip damaged).

DIAGNOSIS. Large bees with dark wings and reticulately rugose propodeum. Male withunique scape extremely dilated, nearly oval (similar to H. satelles; see remarks). Femaleentirely dark, with faint blue reflections on the metasoma.

Distribution. Hawaii; montane wet forest.Male. Measurements: Malar space, 10; eye, 172:172:169:108; clypeus, 102×95; frons,

26×40:110; interalveolar, 31:41:69:77, D 27; interocellar, 38:49:28:49, D 20; scape, 61×90;wing length, 7.4 mm (Hawaii, Kilauea, Auto Road, 4000 ft; BPBM 3447).

Head. Malar space medium-length (ratio about 0.5 DMO). Upper paraocular area andfrons with shallow, round or slightly distorted pits less than 1 pit width apart, pits about thesame size and spacing from the sides to near the middle, ground minutely wrinkled, lustrous;middle with a fine frontal line that may be partly obscured by adjacent punctation and finelongitudinal wrinkles. Lower paraocular depression distinct, extended above antennal socket,


and shiny next to eye. Frontoclypeal sulcus arched. Scape in frontal view extremely dilated,much wider than long, ratio about 1.5; nearly oval in frontal view; frontal surface unevenlyconvex and uniformly covered with fine pits about 1 pit width apart; strongly arched inlateral view with maximum arch near apex of scape; underside with a broad, shallow pit;with medium-length, yellowish white, mostly simple hair. Vertex hair 20-30 or more, white,simple.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground slightly lustrous; hair long(10-22), yellowish white, mostly simple. Mesepisternum with distinct, round pits about 1pit width apart, and long, yellowish white, weakly plumose hair; ground smooth, slightlylustrous. Hypoepimeron punctured, smooth, slightly lustrous; swollen. Metanotum dull.Basal area of propodeum reticulately rugose over entire anterior area and brow, areolaerather large, ground shiny.

Metasoma. Disk of T2 with indistinct, minute, shallow punctation 1-2 pit widths apart,somewhat obscured by smoothly coriaceous, lustrous ground; fine, pale, appressed hair.T7 hair long, dark brown, prostrate.

Terminalia. Lateral process of S7 tapered, length subequal to width of hemisternite.Median process of S8 strongly dilated, dilation membranous, and apex bifurcate with broadtips. Apices of gonoforceps subequal to penis valves.

Color. Head and mesosoma black, metasoma dark brown. Labrum and mandible blackishbrown to black. Face with or without single small mark: clypeus variable, a short triangle,a ragged yellow streak, or no marks; paraocular area and supraclypeal area unmarked;scape unmarked. Pronotum unmarked. Legs reddish brown to black except fore tibia andfemur with obscure pale areas. Wings dark brown.

Female. Head. Malar space medium-length (ratio about 0.6 DMO). Upper paraoculararea and frons with shallow, round or slightly distorted pits less than 1 pit width apart, pitsabout the same size and spacing from the sides to near the middle, ground minutely wrinkled,lustrous; middle with a fine frontal line that may be partly obscured by adjacent punctationand fine longitudinal wrinkles. Apex of clypeus very shallowly emarginate. Dorsal end offacial fovea extended to level of lateral ocellus. Vertex hair 16-30, white, simple.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground slightly lustrous; hair long (12-20), white, simple. Mesepisternum with distinct,round pits about 1 pit width apart, and yellowish white, mostly simple hair; ground smooth,slightly lustrous. Hypoepimeron punctured, smooth, slightly lustrous; swollen. Metanotumdull. Basal area of propodeum reticulate rugose over entire anterior area and brow (rugaeweaker than in male), areolae rather large, ground shiny.

Metasoma. Disk of T2 with shallow, indistinct punctation about 1 pit width apart,somewhat obscured by smoothly coriaceous and lustrous surface; fine, pale, appressedhair, somewhat denser along apical margin in a faint fascia; tergum 1 with faint apical bandof appressed pale hair. T6 hair long, black, erect.

Color. Body and legs dark brown to black except fore tibia with obscure pale area,metasoma with faint metallic blue reflection. Wings dark brown.

Perkins’ collections. Hawaii: Kau, 4000 ft (BMNH); Kilauea (BPBM).Recent collections. Hawaii: Volcano, Old Japanese Schoolhouse, 3750 ft; Puu Puai,

3700 ft.


0.5 mm

A B C D

E F

1.0 mm

Fig. 46. Hylaeus muranus, new combination, male. A. Seventh sternum, ventral view. B. Eighthsternum, ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsalview. E. Face with left scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extentof pubescence shown by dashed lines in B-E.

Flower records. Cheirodendron, Metrosideros.Remarks. Hylaeus muranus, from Hawaii, and H. satelles, from Maui Nui, are sister

taxa and differ in several morphological details. Both are members of the group with darkwings, the metasomal terga finely punctured, and relatively large body size (forewing length:H. muranus, ♂ 6.2-7.4 mm, n = 3; ♀ 7.1-7.8 mm, n = 11; H. satelles, ♂ 6.2-6.9 mm, n =3; ♀ 6.9-7.7 mm, n = 4). Males of the two species have in common a distinct lowerparaocular depression that is shiny along the eye; unmarked legs; a slender, reddish brownmetasoma; and the scape strongly dilated. Hylaeus satelles has the scape slightly less dilatedand circular in profile compared to the much wider and oval profile of the scape of H.muranus. The scape of H. muranus is the most extreme dilation seen among species in theIslands. In both sexes of H. satelles, the propodeum is sparsely sculptured with linear


rugae at the base, if at all, whereas H. muranus is reticulately rugose over most of the basalarea of the propodeum. Based on old specimens, the female of H. satelles has reddishyellow facial marks and light brownish pubescence on the mesosoma. The female of H.muranus has the face black and with pale, whitish pubescence. It also has faint bluishreflections on the metasoma and is somewhat less robust than H. satelles. Both sexes of H.muranus have the upper paraocular area and frons with shallow round pits about 1 pitwidth apart or less, whereas H. satelles has smaller, distorted, denser pits less than 1 pitwidth apart.

References. Perkins, 1907b:97 (record: Hawaii, Kilauea); 1910:604 (key); 1913:82(biogeography).

Hylaeus (Nesoprosopis) mutatus (Perkins)(Fig. 47)

Nesoprosopis mutata Perkins, 1899:93 (♂♀).Hylaeus (Nesoprosopis) mutatus, Meade-Waldo, 1923:34 (list).Nesoprosopis mutata, Daly, 1994:118 (lectotype ♂ BMNH 17a 164; Kauai, Waimea,

2000+ ft).

DIAGNOSIS. Small bees with very short malar spaces and clear to slightly smoky wings.Male face with two or three separate marks; process of S8 undilated and with apex simple,gonoforceps subequal to penis valves. Female face with narrow linear yellow stripes ateach side along eye margin up to the level of the antennal sockets; pronotum sometimeswith large yellow marks on lobes and stripe at each side on collar. The female is similar toH. kauaiensis and H. specularis; it differs from the former in the length of the scutum hairand propodeal sculpture, and from the latter in the punctation of the frons. Hylaeus mutatusalso may have the pronotal lobes marked, which H. kauaiensis never does. The singlefemale H. specularis from Kauai has the lobes unmarked, but females from Hawaii havesuch marks.

Distribution. Kauai; mesic to wet forest.Male. Measurements: Malar space, 3; eye, 110:92:88:57; clypeus, 53×49; frons,

18×24:73; interalveolar, 20:16:45:51, D 18; interocellar, 24:27:20:31, D 14; scape, 35×20;wing length, 3.6 mm (H. mutatus lectotype).

Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular area and fronsuniform to the middle with indistinct, round pits about 1 pit width or less apart, and withminutely wrinkled ground partly obscuring pits, slightly lustrous; faint or no frontal line;small impunctate area above antennal sockets. Lower paraocular depression absent.Frontoclypeal sulcus slightly or not arched. Scape in frontal view variable, weakly tomoderately dilated, width more than half length, ratio about 0.5-0.7; lateral edge in frontalview straight; moderately arched in lateral view with maximum arch near the middle of thescape (contrary to key of Perkins 1910:602); no groove on underside; with medium-length,yellowish white, simple hair. Vertex hair 12-20, yellowish white to pale brown, weaklyplumose.


0.5 mm

A B C D

E F

1.0 mm

Fig. 47. Hylaeus mutatus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view.C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with left scape. F. Scape, lateralview. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shown by dashed lines in B-E.

Mesosoma. Scutum with minute pits 2-3 pit widths apart, ground minutely coriaceous,slightly lustrous; hair short (4-8), yellowish white to pale brown, simple. Metanotum dullto smooth and lustrous. Mesepisternum with distinct, fine, round pits about 2 pit widthsapart, and yellowish white, plumose hair; ground smooth, lustrous. Hypoepimeronimpunctate, smooth, lustrous, slightly swollen. Basal area of propodeum with weak, shortrugae at front, spaced 3-4 rugae apart, otherwise smooth and lustrous to brow.

Metasoma. T2 impunctate; smoothly coriaceous, shiny, with fine, pale, appressed hairsspaced a hair length apart. T7 hair long, dark brown to black, erect.

Terminalia. Lateral process of S7 tapered, length subequal to width of hemisternite.Median process of S8 not or very weakly dilated at base, slightly obtusely angled atmidlength, apex simple and with short pubescence. Apices of gonoforceps equal or shorterthan penis valves.


Color. Labrum and mandible black. Head and mesosoma black; metasoma black to darkbrown. Face with 2 or 3 separate marks: clypeus variable, entirely black or with an irregularsmall subapical spot or with very small subapical spots; lower paraocular areas with triangularmarks almost reaching to antennal sockets or shorter. Pronotum unmarked or with verysmall yellow spot at apex of lobes. Legs black to dark brown, unmarked except fore tibiawith pale area. Wings clear to slightly smoky.

Female. Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular areaand frons uniform to the middle with round pits about 1 pit width apart and with minutelywrinkled ground, slightly lustrous; faint frontal line; small impunctate area above antennalsockets. Apex of clypeus slightly emarginate. Dorsal end of facial fovea extended to levelof lateral ocellus. Vertex hair 10-16, white, weakly plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with minute pits about 2 pitwidths apart, ground minutely coriaceous, slightly lustrous; hair short (2-6), white, simple.Mesepisternum with fine round pits about 2 pit widths apart, and yellowish white, plumosehair; ground smooth, lustrous. Hypoepimeron impunctate, smooth, lustrous, not swollen.Metanotum dull. Basal area of propodeum with weak, short rugae at front, spaced 3-4rugae apart, otherwise smooth and lustrous to brow.

Metasoma. Disk of T2 impunctate; smoothly coriaceous, shiny, with fine, short, pale,appressed hairs spaced more than a hair length apart. T6 hair long, dark brown, erect.

Color. Labrum and mandible black. Face with narrow linear yellow stripes at each sideof lower paraocular areas, extended along eye margin from the level of the antennal socketsto between half and all the way to bottom of eye. Pronotum with or without large yellowmark on lobes and with a stripe at each side on collar. Head and mesosoma black; metasomablack. Legs black, unmarked except fore tibia with pale area. Wings clear.

Perkins’ collections. Kauai: Waimea, 2000 ft (BMNH).Recent collections. Kauai: Awaawapuhi Trail, 3400 ft; Kahuamaa Flat, 3800 ft; Makaha

Rd., 3500 ft; Kuia Valley, 2400 ft; Paaiki Valley, 2700 ft.Flower records. Antidesma platyphyllum, Chamaesyce atrococca, Claoxylon, Styphelia.Remarks. Perkins, in his key (1910:602), placed H. mutatus with species having the

scape flat; however, it is distinctly arched.References. Perkins, 1910:602 (key).

Hylaeus (Nesoprosopis) nalo Magnacca and Daly, new species(Fig. 48)

DIAGNOSIS. Moderately large bees with slightly smoky wings. Head rather large comparedto wing length, with deep paraocular depressions. Median process of S8 very weakly dilated,not swollen; hemisternite of S7 narrow, distinctly longer than wide; apices of gonoforcepsshorter than penis valves, broadly rounded. Female unknown.

Distribution. Oahu; habitat unknown. Known from only one collection.Male. Measurements: Malar space, 8; eye, 156:128:124:88; clypeus, 76×78; frons,



Head. Malar space medium (ratio 0.5 DMO). Upper paraocular area with deep, distinctpits less than 1 pit width apart, smaller and denser on the frons, somewhat distorted nearmiddle; ground lustrous. Lower paraocular depression deep, distinct in frontal view; deepestnear eye. Frontoclypeal sulcus slightly arched. Scape in frontal view moderately dilated,width more than half length, ratio about 0.6; lateral edge in frontal view slightly concave,slightly arched in lateral view; no groove on underside; with medium-length, reddish brownhair. Vertex hair 14-20, pale brown, plumose.

Mesosoma. Scutum with distinct but shallow pits 1-2 pit widths apart, ground minutelycoriaceous, slightly lustrous; hair medium-length to long (12-18), even (varying over area,rather than hairs of different lengths mixed together), golden brown, densely plumose.Scutellum of holotype missing. Mesepisternum with shallow but distinct pits about 1 pit

Fig. 48. Hylaeus nalo, new species, male. A. Seventh sternum, ventral view. B. Eighth sternum,ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E.Face with left scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E.

0.5 mm

A B C D

1.0 mm

E F


width apart or less, and long, brown, plumose hair; ground minutely granular, slightlylustrous. Hypoepimeron shallowly punctate, lustrous; slightly swollen. Metanotum dull.Basal area of propodeum with irregular, widely spaced rugae, smooth coriaceous and shinyin between; granular with close reticular rugae below brow.

Metasoma. Disk of T2 with faintly visible pits and fine, golden, short hairs a hair lengthapart or less. T7 hair short, brown, prostrate.

Terminalia. Lateral process of S7 L-shaped, length distinctly greater than width ofhemisternite. Median process of S8 very weakly dilated, arched, apex bifurcate with long,straplike tips. Apices of gonoforceps slightly shorter than penis valves, rounded.

Color. Head and mesosoma black; metasoma dark reddish brown to black. Labrum andmandible black. Face of type with small, obscure, yellowish brown marks laterad ofsubantennal sulci; probably distinct yellow paraocular marks may be present or absent.Scape unmarked; prothorax unmarked. Legs dark brown to black, unmarked except palearea on fore tibia. Wings slightly smoky.

Female. Unknown.Holotype ♂ labeled “Oahu, Mar. 1914. J. F. Illingworth, collector.” (BPBM).Flower records. None.Etymology. The species epithet nalo is the Hawaiian word meaning lost or forgotten,

referring to the species having languished unrecognized for decades and to the continuinglack of knowledge of its collection site and habitat. Nalo is also the Hawaiian word for awasp or fly; a honey bee is meli nalo.

Remarks. The lack of any other species with deep paraocular depressions on Oahushould make this species immediately identifiable. That together with the weakly dilatedS8 and poorly developed facial marks suggest that it may be related to H. takumiae.Unfortunately the type label gives no specific locality. It most likely inhabits montane areas,but the paraocular depressions, shape of S8, and dark but prostrate T7 hair could indicatea relationship with the coastal H. psammobius.

Hylaeus (Nesoprosopis) niloticus (Warncke), new combination(Fig. 49)

Nesoprosopis obscurata Perkins, 1899:99 (♂♀).Hylaeus (Nesoprosopis) obscuratus, Meade-Waldo, 1923:34 (list).Prosopis nilotica Warncke, 1970:748 (replacement name for Nesoprosopis obscurata

Perkins, junior homonym of Prosopis obscurata Schenck)Nesoprosopis obscurata, Daly, 1994:118 (lectotype ♂ BMNH 17a 151a; Hawaii, Kona;

wing length, 4.8 mm, tip damaged).

DIAGNOSIS. Medium-sized bees with clear to smoky wings. Male scape weakly dilatedand moderately arched; face with one large mark extended at the sides well above antennalsockets as a narrow stripe next to eye, and with supraclypeal area partly yellow or entirelyblack. Female black and unmarked.

Distribution. Hawaii, Lanai, and Molokai; coast and dry lowlands. No recent collections.


Male. Measurements: Malar space, 7; eye, 161:120:110:79; clypeus, 79×72; frons,26×26:102; interalveolar, 15:27:59:69, D 20; interocellar, 36:27:26:31, D 18; scape, 56×24;wing length, 5.0 mm (Molokai, Mts., BPBM 3521).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area with round pitsabout 1 pit width apart; frons with pits closer medially and smaller; ground smooth andlustrous; at middle pits much smaller, contiguous, distorted and separated by a few linearwrinkles, dull; no frontal line evident; very small impunctate area above antennal sockets.Lower paraocular depression very shallow. Frontoclypeal sulcus straight or nearly so. Scapein frontal view weakly dilated, greatest width near apex, width half or less length, ratioabout 0.4; lateral edge in frontal view nearly straight; moderately arched in lateral viewwith maximum arch near middle of scape; with short, yellowish white, mostly simple hair.Vertex hair 10-16, brownish white, weakly plumose.

0.5 mm

A B C D

E F

1.0 mm

Fig. 49. Hylaeus niloticus, new combination, male. A. Seventh sternum, ventral view. B. Eighthsternum, ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsalview. E. Face with left scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extentof pubescence shown by dashed lines in B-E.


Mesosoma. Scutum with pits about 1 pit width apart, ground smooth, lustrous; hairmedium-length (6-14), yellowish white, weakly plumose. Mesepisternum with indistinct,shallow, round pits about 1 pit width apart, and medium-length, yellowish white, plumosehair; ground smoothly coriaceous, lustrous. Hypoepimeron impunctate, smooth, lustrous;slightly swollen. Metanotum dull to slightly lustrous. Basal area of propodeum with widelyspaced, lineate rugae to brow, shiny between rugae; brow weakly rugose, ground nearlysmooth, slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with medium-lengthto short, fine, pale, appressed hairs spaced less than a hair length apart. T7 hairs short,golden, appressed.

Terminalia. Lateral process of S7 tapered, pale, length greater than width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with long, narrow, apically roundedtips. Apices of gonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face with one large mark withsulci and tentorial pits black: clypeus and paraocular area entirely yellow, extended at thesides to well above antennal sockets as a narrow stripe next to eye; supraclypeal area partlyyellow or entirely black; scape unmarked. Pronotum unmarked. Legs brown to black except:fore tibia with pale area, with or without a small, basal yellow spot; fore basitarsus yellowishto brown; mid and hind leg unmarked. Wings clear to slightly smoky.

Female. Head. Malar space medium-length (ratio about 0.6 DMO). Upper paraoculararea with round pits about 1 pit width apart; frons with pits closer medially and smaller;ground smooth and lustrous; at middle pits much smaller, contiguous, but still nearly round,lustrous; faint frontal line evident; no impunctate area above antennal sockets. Apex ofclypeus shallowly emarginate. Dorsal end of facial fovea extended to level of lateral ocellus.Vertex hair 6-10(-16), yellowish white, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with pits about 1 pit widthapart, ground smooth, lustrous; hair short (4-6), white, simple. Mesepisternum withindistinct, shallow, round pits about 1 pit width apart, and short, yellowish white, plumosehair; ground smoothly coriaceous, lustrous. Hypoepimeron impunctate, smooth, lustrous;slightly swollen. Metanotum dull to slightly lustrous. Basal area of propodeum with irregularlineate rugae to brow, spaced 2-4 rugae apart, surface shiny between rugae; brow weaklyrugose, ground nearly smooth, slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with short, fine, pale,appressed hairs spaced about one hair length apart. T6 hair long, golden, appressed.

Color. Head and mesosoma black; legs and metasoma dark brown; unmarked. Wingssmoky.

Perkins’ collections. Hawaii: Kona (BMNH, OUMNH), coast (BPBM). Molokai:Molokai Mts. (BMNH, BPBM), 2000 ft (BPBM).

Recent collections. None.Flower records. None.Remarks. Colors of integument and pubescence are based on old specimens. The new

name Prosopis nilotica was proposed by Warncke (1970), who considered Prosopis to be


the correct name of the genus; his reason for naming a Hawaiian species “nilotica” (i.e., ofthe Nile) is unclear. Warncke also proposed the new name P. murana for H. insignis (bothare homonyms of names that Meade-Waldo [1923] considered synonyms of other species),as well as unwarranted names for H. laticeps, H. nivalis, and H. vicinus, homonyms alreadycorrected by Meade-Waldo (1923). See remarks for H. assimulans.

References. Perkins, 1910:602 (key); 1911:727 (Blackburn had series of mixed species).Fullaway, 1918:396 (record: Hawaii, Kawaihae, Kilauea). Suehiro, 1986:41, 60 (record:Hawaii, Kona).

Hylaeus (Nesoprosopis) nivicola Meade-Waldo(Fig. 50)

Nesoprosopis nivalis Perkins, 1899:83-84 (♂♀; Pl. 2, Fig. 13; flower: Styphelia).Hylaeus (Nesoprosopis) nivicola Meade-Waldo, 1923:34 (list; replacement name for

Nesoprosopis nivalis Perkins, a junior homonym of Hylaeus nivalis Morawitz).Prosopis farinosa Warncke, 1970:748 (unwarranted replacement name for Nesoprosopis

nivalis Perkins) (new synonymy)Nesoprosopis nivalis, Daly, 1994:118 (lectotype ♂ BMNH 17a 136a; Maui, Haleakala,


DIAGNOSIS. Medium-sized bees with wings clear to slightly smoky and with uniqueextremely long malar spaces. Male with unique ridge just below antennal sockets; facewith clypeus and supraclypeal area yellow and with small yellow marks in adjacent paraocularareas. Female black and unmarked. See remarks.

Distribution. Maui; occurring only around Haleakala crater, common in alpine andsubalpine dry shrubland down to 6000 ft.

Male. Measurements: Malar space, 18; eye, 138:128:108:87; clypeus, 72×69; frons,28×26:102; interalveolar, 14:26:59:69, D 23; interocellar, 31:36:27:36, D 15; scape, 46×34;wing length, 4.9 mm (Maui, Haleakala, 9000 ft, BPBM 3520).

Head. Malar space extremely long (ratio about 1.2 DMO). Upper paraocular area withfine pits less than 1 pit width apart, crowded into subpolygonal shapes and more or lessaligned in concentric rows separated by minute wrinkles, slightly lustrous; below puncturedarea and along the eye are dense, minute, parallel wrinkles; frons dull, with pits smaller anddenser along middle; fine frontal line may be evident, with a minute, central, elliptical holein some bees; median, impunctate, slightly lustrous area above antennal sockets. Lowerparaocular depression distinct, deep along eye, with dense, minute, parallel wrinkles andbordered by an elevated, rounded ridge beneath antennal socket. Frontoclypeal sulcus arched.Scape in frontal view moderately dilated, width more than half length, ratio about 0.7;lateral edge in frontal view distinctly rounded (convex); moderately arched in lateral viewwith maximum arch near middle or slightly toward apex of scape; no groove on underside;with long, yellowish white, simple hair. Vertex hair 20-30 or more, pale brown, simple.

Mesosoma. Scutum with fine pits about 1 pit width apart, ground minutely wrinkled,dull to slightly lustrous; hair long (14-30), yellowish white to pale brown, plumose.Mesepisternum with long, white, plumose hair; pits variable, indistinct or relatively distinct,


round, shallow, 1 pit width apart or less; ground variable, minutely roughened, wrinkled orstriate, dull to slightly lustrous. Hypoepimeron impunctate, smooth to striate. Metanotumdull. Basal area of propodeum lineately rugose at the front, shiny between rugae, becomingreticulate near brow; brow granularly coriaceous, dull to slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous to shiny, with veryfine, short, pale hair more than a hair length apart. T7 hair medium-length, golden, appressed.

Terminalia. Lateral process of S7 tapered, length subequal to width of hemisternite.Median process of S8 linear and with apex simple or nearly so, not distinctly bifurcate.Apices of gonoforceps longer than penis valves, but not pointed or strongly curved inward.

Color. Body black. Labrum and mandible blackish brown to black. Face with clypeusand supraclypeal area entirely yellow or nearly so; lower paraocular area variable, stripe of

Fig. 50. Hylaeus nivicola, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.

0.5 mm

A B C D

E F

1.0 mm


yellow below antennal socket and along sulcus (at least partly separated from clypeal markby narrow black line) or area unmarked; scape unmarked. Pronotum unmarked. Fore legdark brown to black except tibia with pale area and basal yellow spot. Mid leg dark brownto black except tibia unmarked or with obscure or distinct small yellow spot at very base,basitarsus variable, with or without some yellow marking. Hind leg dark brown to blackexcept tibia with yellow stripe in basal third to half, basitarsus with yellow marking onbasal half or more. Wings clear to slightly smoky.

Female. Head. Malar space extremely long (ratio about 1.1 DMO). Upper paraoculararea and frons with fine pits less than 1 pit width apart, crowded into subpolygonal shapesand more or less aligned in concentric rows separated by minute wrinkles; fine frontal linein some bees. Apex of clypeus shallowly emarginate. Dorsal end of facial fovea extendedto level of lateral ocellus. Vertex hair 16-24, pale brown, simple.

Mesosoma. Fore tarsus with hair apically curved. Scutum with pits 1-2 pit widths apart,ground slightly lustrous; hair long (12-30), pale brown, plumose. Scutellum of some beeswith pits more than 2 pit widths apart. Mesepisternum with pits relatively distinct comparedto male, round, shallow, 1 pit width apart or less, with long, white, plumose hair; groundvariable, minutely wrinkled or striate, dull to slightly lustrous. Hypoepimeron impunctate,smooth to striate. Metanotum dull. Basal area of propodeum lineately rugose at front,often long, parallel rugae, spaced close together in some specimens, shiny between rugae,becoming weaker and reticulate near or beyond brow; brow granularly coriaceous, slightlylustrous to dull.

Metasoma. Disk of T2 impunctate, lustrous to polished, with bare areas or extremelyshort, very fine, appressed hair. T6 hair medium-length, pale brown, appressed.

Color. Body and legs black; unmarked. Wings clear.Perkins’ collections. Maui: Haleakala, 6000 ft (BMNH, BPBM), >6000 ft (BPBM),

7000 ft (BPBM), 7-10,000 ft (BPBM), 9000 ft (BMNH, BPBM), 9 to 10,000 ft (BMNH),crater, 8000 ft (OUMNH).

Recent collections. Maui: Haleakala Science City, 3010 m; near White Hill, 2860 m;Kalahaku, 2835 m; Kalepeamoa, 2835 m; Kuiki, 2170 m; Leleiwi, 2710 m; Haleakala N. P.Service Area, 2073 m; Laie flats, 2060 m; Lauulu Tr., 2015 m; near Kapalaoa Cabin, 2195m; Sliding Sands Tr., 9000 ft; Halemauu Tr., 7800 ft; Puu Keokea, 7100 ft; Koolau ForestReserve, 2158 m.

Flower records. Argyroxiphium sandwicense, Coprosma montana, Dodonaea, Dubautiamenziesii, Geranium cuneatum, Oenothera sp., Sophora, Styphelia.

Remarks. Hylaeus nivicola, from the high slopes and crater of Maui’s Haleakala Volcano,and H. volcanicus, from Hawaii and Maui, are sister taxa. They are sympatric in the craterof Haleakala. Both sexes have in common malar spaces longer than those of the allied H.laetus and H. difficilis (see also remarks for H. difficilis); and both have the similar lineatelyrugose anterior area of the propodeum. Males have in common the depressed paraoculararea and similar facial marks with conspicuous black tentorial pits (paraocular area lessdepressed and tentorial pits inconspicuous in the allied H. laetus and H. difficilis). Malesof H. nivicola are distinguished by the much longer malar space (the longest of any speciesin the Islands), rounded ridge beneath the antennal socket (absent in H. volcanicus), morestrongly dilated scape with convex lateral margin in frontal view (only slightly dilated and


with the margin slightly concave in H. volcanicus), and the simple apex of the medianprocess of S8 (bifurcate with long filamentous tips in H. volcanicus). Females of H. nivicolahave a longer malar space, more densely punctured frons with subpolygonal pits, and reddishyellow hair at the tip of the metasoma (malar space shorter; frons with mostly round pits;pale, yellowish hair at the apex of the metasoma in H. volcanicus).

References. Perkins, 1910:601 (key); 1913:80 (flower: Styphelia, referred to asCyathodes). Swezey, 1936:139 (flower: silversword; record: Maui, Haleakala Crater).Beardsley, 1980:40 (record: Maui, Haleakala).

Hylaeus (Nesoprosopis) ombrias (Perkins)(Fig. 51)

Nesoprosopis ombrias Perkins, 1910:604-605 (♂♀; key).Hylaeus (Nesoprosopis) ombrias, Meade-Waldo, 1923:35 (list).Nesoprosopis ombrias, Daly, 1994:118 (lectotype ♂ BPBM; Hawaii, S. Kona, 3000 ft;


DIAGNOSIS. Large bees with dark smoky wings and terga without punctation (unusualfor bees of this size). Male with unusual scape, slightly dilated, widest near the middle andsmaller at both ends, and with short, plumose hair along the median edge; face with two orthree separate yellow marks (in paraocular areas and with or without a mark on the clypeus,no supraclypeal mark), otherwise body and legs black and unmarked; distinguished fromthe smaller, less melanic, sister species, H. assimulans, by reduced facial marks (none onsupraclypeal area, sometimes none on clypeus), lineate to reticulate rugosity often to browof propodeum, and darker wings. Female black and unmarked, upper frons granular.

Distribution. Hawaii; coast and montane dry shrubland and forest.Male. Measurements: Malar space, 5; eye, 169:137:128:92; clypeus, 84×84; frons,

28×31:109; interalveolar, 18:31:61:72, D 26; interocellar, 36:36:23:38, D 18; scape, 61×29;wing length, 6.3 mm (Hawaii, S. Kona, 3000 ft, BPBM 3525).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area and frons withround pits less than 1 pit width apart; ground minutely wrinkled, dull; a small, dark, medianarea of smaller, much denser pits; shiny area below dark spot and above antennal sockets.Lower paraocular depression very shallow. Frontoclypeal sulcus straight to slightly arched.Scape in frontal view slightly dilated and widest at middle, width half length, ratio about0.5 ; lateral edge in frontal view nearly straight; moderately arched in lateral view withmaximum arch near middle of scape; no groove on underside; with short, brownish white,plumose hair. Vertex hair 10-18, brown, plumose.

Mesosoma. Scutum with shallow, indistinct pits 1-2 pit widths apart, ground lustrous;hair medium-length (8-12), yellowish white, plumose. Scutellum with pits distinct andlarger, ground lustrous. Mesepisternum with barely discernible to indistinct pits, about 1pit width apart, and medium-length, yellowish white, plumose hair; ground minutelycoriaceous, slightly dull. Hypoepimeron impunctate, slightly lustrous, slightly swollen.


0.5 mm

A B C D

E F

1.0 mm

Fig. 51. Hylaeus ombrias, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.

Metanotum reticulately rugose, slightly lustrous. Basal area of propodeum with largerlineate and smaller reticulate rugae to brow; brow granularly coriaceous and slightly lustrous.

Metasoma. Disk of T2 impunctate (unusual for bee of this large size), smoothlycoriaceous, and lustrous; fine, pale, appressed hairs spaced about one hair length apart. T7hair medium-length, golden, appressed.

Terminalia. Lateral process of S7 tapered, length greater than width of hemisternite.Median process of S8 weakly dilated at flexure with a slight peak, apex bifurcate withnarrow straplike tips. Apices of gonoforceps subequal to penis valves.

Color. Head and mesosoma black; metasoma dark brown. Labrum and mandible blackishbrown to black. Face with two or three separate marks: clypeus with or without a yellowmark, supraclypeal area unmarked; lower paraocular area with semi-triangular yellow mark


along the eye up to or above level of antennal socket; scape unmarked. Pronotum unmarked.Legs dark brown to black; unmarked except fore tibia with pale area. Wings dark smoky.

Female. Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area withdistinct, round pits about 1 pit width or more apart, ground minutely wrinkled along eye,face slightly dull; frons with similar pits round and uncrowded, slightly smaller mediallyand with ground minutely wrinkled; at middle, pits may or may not be obscured by minutelywrinkled ground parallel to midline; frontal line distinct. Apex of clypeus distinctlyemarginate. Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 6-10(up to 20 in some montane specimens), brown, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with poorly defined minutepits, 1-2 pit widths apart and obscured by sculpture of ground; hair white to pale brown,short (4-6), plumose. Mesepisternum with barely discernible to indistinct pits, about 1 pitwidth apart, and short, brownish to white, weakly plumose hair; ground minutely coriaceousto strongly striate, slightly dull. Hypoepimeron impunctate, slightly lustrous, slightly swollen.Metanotum reticulately rugose, slightly lustrous. Basal area of propodeum with sculpturevariable, reticulate or lineately rugose to brow.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with short, fine, pale,appressed hairs spaced about one hair length apart. T6 hair medium-length, brown,appressed.

Color. Body black. Legs black; unmarked except fore tibia with pale area. Face andpronotum unmarked. Wings dark smoky.

Perkins’ collections. Hawaii: Kona, S. (OUMNH), 1800 ft, 2000 ft, 3000 ft (BPBM).Recent collections. Hawaii: Ahumoa, 5900 ft; Kipuka Kalawamana, 5000 ft; Puu

Waawaa, 4200 ft; South Point, coast.Flower records. Bidens menziesii, Chamaesyce olowaluana, Scaevola sericea, Sida,

Tribulus cistoides.Remarks. See remarks for H. assimulans and H. flavipes.References. Suehiro, 1986:60 (record: Hawaii, Kona).

Hylaeus (Nesoprosopis) paradoxicus (Perkins)(Fig. 52)

Nesoprosopis paradoxica Perkins, 1899:111-112 (♂♀; Pl. 2, Figs. 7, 25; nest: Sophora).Hylaeus (Nesoprosopis) paradoxicus, Meade-Waldo, 1923:35 (list).Nesoprosopis paradoxica, Daly, 1994:118 (lectotype ♂ BMNH 17a 141b; Hawaii, Kona,

4000 ft; wing length, 8.0 mm).Nesoprosopis erythrodemas Perkins, 1899:112 (♀; holotype ♀ BMNH 17a 147; Hawaii,

Kau, 4000 ft; wing length, 7.8 mm) (new synonymy).Hylaeus (Nesoprosopis) erythrodemas, Meade-Waldo, 1923:35 (list).Nesoprosopis erythrodemas, Daly, 1994:117 (holotype confirmed).

DIAGNOSIS. Large bees with punctured terga, dark wings, and unique body color: headand mesosoma black and metasoma red. Sister species of H. gliddenae; see remarks forthat species.


Distribution. Hawaii; montane dry and mesic forest.Male. Measurements: Malar space, 13; eye, 187:159:156:113; clypeus, 110×102; frons,

33×38:115; interalveolar, 26:38:61:72, D 28; interocellar, 40 44:36:49, D 20; scape, 63×54;wing length, 8.0 mm (Hawaii, Kona, 4000 ft, BPBM 3528).

Head. Malar space medium-length (ratio about 0.6 DMO). Upper paraocular area withindistinct, shallow pits about 1 pit width apart; frons slightly concave, with pits smallermedially, contiguous, distorted, and near the middle pits obliterated by fine, parallel,longitudinal wrinkles; impunctate, lustrous area above antennal sockets with mediandepression or shallow hole at middle. Lower paraocular depression shallow. Frontoclypealsulcus slightly arched or nearly straight. Scape in frontal view moderately dilated, widthmore than half length, ratio about 0.8; lateral edge in frontal view distinctly convex; strongly

0.5 mm

B C DA

E F

1.0 mm

Fig. 52. Hylaeus paradoxicus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


arched in lateral view with maximum arch near middle of scape; no groove on underside;with medium-length, yellowish white, mixed simple and weakly plumose hair. Vertex hair20-30, white, weakly plumose.

Mesosoma. Scutum with distinct pits 1-2 pit widths apart, ground slightly granularlycoriaceous, slightly lustrous; hair medium-length to long (10-20), yellowish white, weaklyplumose. Scutellum with slightly larger pits, smoothly coriaceous, lustrous. Mesepisternumwith distinct, round pits about 1 pit width apart, and medium-length, yellowish white,plumose hair; ground smooth, lustrous. Hypoepimeron punctured, smooth, lustrous, swollen.Metanotum dull. Basal area of propodeum protuberant in middle, roughened overall byreticulate rugae except the posterior, vertical, coriaceous face; rugae and ground granularor minutely wrinkled, dull to slightly lustrous; lateral carina strong, terminating anterolaterallyin a protuberant area.

Metasoma. Disk of T2 with distinct punctation, minute, shallow pits about 2 pit widthsapart; ground smoothly coriaceous, lustrous; with fine, pale, short hair. T7 hair long, black,erect.

Terminalia. Lateral process of S7 tapered, with length less than width of hemisternite.Median process of S8 linear, apex bifurcate with rather long, straplike tips. Apices ofgonoforceps longer than penis valves; gonoforceps rather narrow and with long, apical,plumose hair.

Color. Labrum and mandible blackish brown to black. Face unmarked. Scape in frontalview with obscure yellow stripe on lateral margin. Pronotum unmarked or with well-separated lateral marks on collar. Head and mesosoma black; metasoma red; T2 with anelongate black spot laterally behind spiracle. Legs dark brown to black; unmarked exceptpale area on fore tibia, all tarsi with or without reddish markings. Wings dark smoky.

Female. Head. Malar space medium-length (ratio about 0.6 DMO). Upper paraoculararea with distinct, round pits about 1 pit width or more apart, ground smoothly coriaceousand shiny; frons with pits smaller medially, but still round and uncrowded, ground minutelywrinkled and lustrous; middle from median ocellus to acme depressed about midway, andat this point the fine frontal line distinct and wide to acme. Apex of clypeus shallowlyemarginate. Mandible weakly three-toothed, the two anterior (dorsal) teeth poorly separated.Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 20-30, white,weakly plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground smoothly coriaceous, lustrous; hair medium-length to long (10-20), yellowishwhite, weakly plumose. Scutellum similar, with slightly larger pits. Mesepisternum withdistinct, round pits about 1 pit width apart, and medium-length, yellowish white, plumosehair, dense on sides of propodeum; ground smooth, lustrous. Hypoepimeron punctured,smooth, lustrous, swollen. Metanotum dull. Basal area of propodeum peculiarly shapedwith somewhat protuberant, short, anterior area bearing rather weak, widely spaced, lineaterugae to brow; rugae, spaces between rugae, and ground below brow slightly granularlycoriaceous, slightly lustrous; weak lateral carinulae terminating anterolaterally in aprotuberant area.

Metasoma. Disk of T2 with distinct punctation; minute, shallow pits about 2 or morepit widths apart; ground smoothly coriaceous, shiny; with fine, pale, short hair; T6 hairdark reddish brown to black.


Color. Head and mesosoma black; metasoma red. Legs dark brown; unmarked exceptfore tibia with pale area. Face and pronotum unmarked. Wings dark smoky.

Perkins’ collections. Nesoprosopis paradoxica: Hawaii: Kilauea (BPBM); Kona, 4000ft (BMNH, BPBM, OUMNH); Kona, N., 3-4000 ft (BPBM); Kona, S., 2000 ft (BPBM).N. erythrodemas: Hawaii: Kau, 4000 ft (BMNH); Kilauea (BPBM); 4000 ft (BPBM,OUMNH).

Recent collections. Hawaii: Puu Waawaa, 4100 ft; Kipuka Alala, 5800 ft.Flower records. Chamaesyce olowaluana, Myoporum.Remarks. We conclude that H. erythrodemas is a rare, shiny red, female variant and a

synonym of H. paradoxicus. We chose the latter name because it is based on a male type.Female H. erythrodemas are nearly identical to female H. paradoxicus. Examination of theholotype female of H. erythrodemas confirms some of the differences mentioned by Perkins:body less robust, propodeum with much weaker rugosity, metasoma red, second tergumsparsely punctured (also much smaller pits), and second sternum strongly raised. The shapeof the face and propodeum, however, appear the same as H. paradoxicus, and otherspecimens identified as H. erythrodemas are less distinctly different in the characters listedby Perkins. Both nominal taxa are known only from Hawaii: H. erythrodemas was describedfrom Kau at 4000 ft and H. paradoxicus from Kona at 4000 ft. Hylaeus erythrodemas isknown only from the female, whereas both males and females are known for H. paradoxicus.Specimens identified by Perkins as H. erythrodemas are: 3♀ Kilauea (BPBM); 1♀ Kilauea,4000 ft (OUMNH); ♀ holotype, Kau, 4000 ft (BMNH). Specimens available of H.paradoxicus are: many ♂♂ and a few ♀♀ from Kilauea, 4000 ft; Kilauea Dry Forest, 4000ft; Kona, 4000 ft; N. Kona, 3000 ft; and S. Kona, 2000 ft (BPBM); 1♂ 1♀ Kona, 4000 ft(OUMNH); ♂ lectotype, Kona, 4000 ft (BMNH). It is evident that specimens identified asH. paradoxicus are widespread and sympatric with those identified as H. erythrodemas.All recent specimens have come from Kona.

References. Perkins, 1910:604 (key); 1913:79 (morphology). Suehiro, 1986:61 (record:Hawaii, Kona). Hylaeus erythrodemas: Perkins, 1907b: 97 (record: Hawaii, Kilauea).

Hylaeus (Nesoprosopis) pele (Perkins)(Fig. 53)

Nesoprosopis pele Perkins, 1911:723-724 (♂; Hawaii, Kilauea).Hylaeus (Nesoprosopis) pele, Meade-Waldo, 1923:35 (list).Nesoprosopis pele, Daly, 1994:116 (type not located).

DIAGNOSIS. Small bees with clear to slightly smoky wings. Male face with three large,but separate marks, almost filling the clypeus and the lower paraocular areas below theantennal sockets; scape moderately dilated and with long hair along median edge; fronswith indistinct, small patch of pale hair; legs and pronotal lobes may be marked with yellow,but not the pronotal collar. Female face unmarked or with short, irregular yellow marksalong eyes; mid and hind tibia with or without yellow marks at bases; pronotum black.

Distribution. Hawaii; dry shrubland and dry and mesic forest.


Male. Measurements: Malar space, 3; eye, 86:83:76:51; clypeus, 47×42; frons, 15×20:64;interalveolar, 10:17:39:47, D 12; interocellar, 20:27:15:25, D 11; scape, 29×22; wing length,3.3 mm (Hawaii, Kipuka Alala, 5800 ft, KM-99-141).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area and fronssomewhat convex, paraocular area with nearly round pits about 1-2 pit widths apart ormore, on frons pits closer to about 1 pit width apart, at middle pits contiguous and slightlydistorted; ground smoothly coriaceous, shiny, especially where pits are sparse below ocelli,less so in paraocular area above antennal sockets; small patch of erect, pale hairs at middle;impunctate, lustrous area above antennal sockets. Frontal line absent. Lower paraoculararea with slight depression. Frontoclypeal sulcus slightly arched. Scape in frontal viewmoderately dilated, width more than half length, ratio about 0.8; lateral edge in frontalview nearly straight; moderately arched in lateral view with maximum arch near middle ofscape; underside of median edge with a groove; with long, simple, yellowish white hair.Vertex hair 6-16, yellowish white, simple.

Mesosoma. Scutum nearly impunctate, only minute shallow pits visible in certain lights,2-4 pit widths apart, pits obscured by granular ground, dull in comparison with back ofhead and brow of propodeum; although pits are obscure, the cuticular surface of scutum atsides when seen in certain lights is uneven, with small, faint depressions associated withpits; hair medium-length (6-12), yellowish white, weakly plumose. Mesepisternum virtuallyimpunctate, only minute pits visible 2-4 pit widths apart and with long, yellowish white,plumose hair; ground smoothly coriaceous, lustrous. Hypoepimeron impunctate, smooth,lustrous, swollen. Metanotum finely wrinkled, dull to slightly lustrous. Basal area ofpropodeum minutely wrinkled or roughened at basal quarter to half, becoming smoothlycoriaceous and shiny at brow; brow prominent.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, shiny; with short, fine, pale,appressed hairs spaced more than a hair length apart or more. T7 hair long, red-brown,erect.

Terminalia. Lateral process of S7 tapered, narrow, length greater than width ofhemisternite. Median process of S8 moderately dilated, apex bifurcate with moderatelylong, rounded tips, about half as broad as long. Apices of gonoforceps longer than penisvalves.

Color. Labrum and mandible black. Face with three separate yellow marks: clypeus notquite filled with an irregular mark, leaving black margin; lower paraocular area with markup to level of antennal socket; supraclypeal area and scape unmarked. Highly markedspecimens may have the clypeus almost entirely yellow and paraocular marks extendedslightly above antennal sockets. Pronotum unmarked or with very small spots on lobes.Legs dark brown to black except: fore tibia and apex of fore femur with pale areas that maybe yellow along dorsal margins in some specimens, fore basitarsus yellow; mid tibia with orwithout small yellow spots at base and apex, mid basitarsus yellow to partly yellow; hindtibia with or without yellow mark in basal third, hind basitarsus yellow to black. Wingsslightly smoky to clear and iridescent.

Female. Head. Malar space medium-length (ratio about 0.5 DMO). Upper paraoculararea and frons somewhat convex, with nearly round pits about 1-2 pit widths apart, ground


smoothly coriaceous, lustrous; at middle pits obscured by minutely wrinkled ground alongfaint frontal line; small impunctate, minutely wrinkled area above antennal sockets. Apexof clypeus shallowly emarginate. Dorsal end of facial fovea not or barely extended to levelof lateral ocellus. Vertex hair 6-10, yellowish white, simple.

Mesosoma. Fore tarsus with hair apically curved. Scutum virtually impunctate, onlyminute pits visible, 2-4 pit widths apart, obscured by granular ground, slightly lustrous(dull in comparison with top of head and brow of propodeum); hair short (2-6, few longerto 10), white. Mesepisternum virtually impunctate, only minute pits visible, 2-4 pit widthsapart and with long, yellowish white, plumose hair; ground smoothly coriaceous, lustrous.Hypoepimeron impunctate, smooth, lustrous, slightly swollen. Metanotum finely wrinkled,dull to slightly lustrous. Basal area of propodeum minutely wrinkled or roughened at basal

0.5 mm

A B C D

E F

1.0 mm

Fig. 53. Hylaeus pele, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view, andapex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with left scape.F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shown bydashed lines in B-E.


half, becoming smooth and shiny at brow, or basal area almost all smooth; brow prominent.Dorsum of mesosoma slightly depressed, almost flat in lateral view.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, shiny; with short, fine, pale,appressed hairs spaced more than a hair length apart. T6 hair long, light brown, erect.

Color. Body black and legs blackish brown to black except: lower paraocular area withor without short, irregular yellow mark along eye; fore tibia and apex fore femur with paleareas; mid and hind tibia with or without yellow mark at base. Wings clear, iridescent.

Perkins’ collections. None. Type collected later by Perkins at Kilauea, Hawaii.Recent collections. Hawaii: Kipuka Nene, 2900 ft; Kipuka Puaulu, 3900 ft; Mauna

Loa Rd., 4100 ft; Puu Waawaa, 4100 ft; Kipuka Alala, 5700 ft; Ahumoa, 5900 ft; PuuHuluhulu (N. Hilo), 6700 ft.

Flower records. Bidens menziesii, Chamaesyce olowaluana, Crocosmia ×crocosmiiflora*, Myoporum, Sophora.

Remarks. The type of H. pele, taken at Kilauea, has not been located. Males collectedat Kilauea and Kona generally match the Latin description, some with more yellow on theface and legs than Perkins describes (as well as sometimes marks on the pronotal lobes).The species is relatively common throughout the drier upper slopes of Mauna Loa andMauna Kea as well as at Kipuka Nene.

References. None.

Hylaeus (Nesoprosopis) perkinsianus (Timberlake)(Fig. 54)

Nesoprosopis perkinsiana Timberlake, 1926:22-23 (♂♀; holotype ♂ BPBM 226; Nihoa,summit 856 ft, flower: probably Sida; length of bodies of 4♂ was 6-7 mm).

DIAGNOSIS. Medium-sized bees with clear wings. Male face with yellow mark (clypeusand supraclypeal area, with or without mark in lower paraocular areas) divided by fineblack sulci, unusual lateral process of S7 very short, median process of S8 not dilated andapex simple; unusual gonoforceps expanded apically beyond penis valves. Female blackand unmarked. Sister species of H. hirsutulus; see remarks.

Distribution. Nihoa; probably occurs over the entire island. Timberlake (1926:23) andBeardsley (1966:166) records; no recent collections.

Male. Measurements: Malar space, 5; eye, 142:120:115:79; clypeus, 74×74; frons,19×28:95; interalveolar, 19:27:55:64, D 20; interocellar, 32:32:26:33, D 18; scape, 54×25;wing length, 5.2 mm (Nihoa Island, BPBM 3531).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area and frons withround, shallow pits less than 1 pit width apart; ground minutely wrinkled, slightly lustrous;median, impunctate, lustrous area just above antennal sockets; frontal line evident. Lowerparaocular depression distinct, but shallow. Frontoclypeal sulcus straight. Scape in frontalview slightly dilated, width equal to half length, ratio about 0.5; lateral edge in frontal viewconcave; moderately arched in lateral view with maximum arch near middle of scape; no


groove on underside; with medium, brownish white, plumose hair. Vertex hair long, palebrown, plumose.

Mesosoma. Scutum with round pits about 2 pit widths apart, ground smoothly coriaceous,slightly lustrous; hair short (6-10), brownish white, plumose. Scutellum with pits slightlylarger, smoothly coriaceous, lustrous. Mesepisternum with distinct, round pits, about 1 pitwidth apart, and long, yellowish white, weakly plumose hair; ground minutely coriaceousto striate, slightly lustrous. Hypoepimeron mostly impunctate, smooth, slightly lustrous,slightly swollen. Metanotum dull. Basal area of propodeum lineately rugose to brow, shinybetween rugae at front; brow granularly coriaceous, slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with short, fine, pale,appressed hairs spaced more than a hair length apart. T7 hair short, light brown, prostrate.

Fig. 54. Hylaeus perkinsianus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-D.

0.5 mm

A B C D

E F


Terminalia. Lateral process of S7 very short, blunt, with length less than width ofhemisternite. Median process of S8 linear and with apex simple. Apices of gonoforcepsexceeding penis valves.

Color. Labrum and mandible blackish brown to black. Face with one large mark withfine black sulci in “H” shape: clypeus entirely yellow or nearly so; lower paraocular areawith or without a narrow yellow stripe along sulci and not extended above antennal sockets;supraclypeal area to acme yellow; scape unmarked. Pronotum unmarked. Legs dark brownto black and unmarked except fore tibia with pale area and hind tibia with or withoutobscure basal spot. Wings nearly clear.

Female. Not examined; according to Timberlake (1926:23) the female is very similar tothe female of H. hirsutulus, except that the wings are subhyaline instead of fuscous.

Perkins’ collections. None.Recent collections. Beardsley (1966:166) from Nihoa.Flower records. Sida.Remarks. Timberlake (1926:22) stated that H. perkinsianus was closely allied to H.

hirsutulus from Kauai, a conclusion supported here. The males of the two forms are verysimilar (short to very short malar space, wide face, similar facial marks, scape slightlydilated and moderately arched, linear process of S8 with simple apex) and share the veryshort lateral processes on S7 not found in other species. Timberlake observed differencesin the configuration of the apices of the gonoforceps: narrower and rounded apically in H.perkinsianus; broader and slightly angular apically in H. hirsutulus. The legs of H.perkinsianus are unmarked or with only obscure yellow spots at the tibial bases and thewings nearly clear, whereas H. hirsutulus has at least a large, basal yellow mark on the hindtibia and dark smoky wings. We confirm Timberlake’s observations and treat H. perkinsianusas a distinct species until additional studies prove otherwise.

References. Bryan, 1926:9 (record: Nihoa). Williams, 1927:429 (record: Nihoa).Beardsley, 1966:166 (record: Nihoa). Daly, 1994:120 (list).

Hylaeus (Nesoprosopis) perspicuus (Perkins)(Fig. 55)

Nesoprosopis perspicua Perkins, 1899:109 (♂♀; Pl. 2, Fig. 5; flower record: Straussia).Hylaeus (Nesoprosopis) perspicuus, Meade-Waldo, 1923:35 (list).Nesoprosopis perspicua, Daly, 1994:118 (lectotype ♂ BMNH 17a 143b; Kauai,

Makaweli, 2000 ft).

DIAGNOSIS. Large bees with punctured terga, dark wings, tibiae mostly white, and witha unique broad, white pronotal collar.

Distribution. Kauai; montane mesic and wet forest. No recent collections.Male. Measurements: Malar space, 6; eye, 179:151:151:99; clypeus, 92×86; frons,

28×40:111; interalveolar, 28:32:65:74, D 26; interocellar, 34:40:34:43, D 20; scape, 55×37;wing length, 6.3 mm (H. perspicuus lectotype).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area at side with pitsless than 1 pit width apart; across frons to middle with pits medially smaller, contiguous,


distorted, subpolygonal; ground obscured by punctation; small impunctate area aboveantennal socket. No frontal line. Upper face with long, yellowish white, plumose hair.Lower paraocular depression absent. Frontoclypeal sulcus slightly arched. Scape in frontalview moderately dilated, width more than half length, ratio about 0.7; lateral edge in frontalview nearly straight; moderately arched in lateral view with maximum arch near middle ofscape.

Mesosoma. Scutum with distinct pits about 1 pit width apart, ground nearly smooth,lustrous. Scutellum with slightly larger pits, smooth, lustrous. Mesepisternum with distinct,shallow pits less than 1 pit width apart, and light brownish white, weakly plumose hair;ground nearly smooth, lustrous. Hypoepimeron obscurely punctured in lower half, smooth,lustrous, swollen. Basal area of propodeum convex in middle, weakly reticulately rugosenearly to brow area, shiny between rugae; brow indistinct, nearly smooth, lustrous. Lateralcarina without an anterolateral protuberance.

Metasoma. Disk of T2 with punctation; smoothly coriaceous, lustrous; with bare areasotherwise with fine, pale, appressed hairs spaced about one hair length apart.

Terminalia. Lateral process of S7 parallel-sided (not dissected and partly hidden fromview), with length probably greater than width of hemisternite. Median process of S8moderately dilated, apex bifurcate with moderately long, straplike, rounded tips. Apices ofgonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Head and mesosoma dark blackishbrown; metasoma dark reddish brown. Face unmarked except scape with lateral marginyellowish. Pronotum with collar exceptionally broad, entirely ivory; lobes dark. Legs reddishbrown, fore tibia with pale area, fore and mid tibiae marked full length with ivory stripe,hind tibia ivory, except in caudal view with dark spot in apical half. Wings dark smoky.

Female. Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area andfrons with shallow, slightly distorted pits, less than 1 pit width apart and more or less

Fig. 55. Hylaeus perspicuus, male. A. Face with left scape. B. Scape, lateral view.

A B


aligned in rows separated by fine, parallel wrinkles (wrinkles stronger or weaker in differentbees); at the sides the wrinkles curve broadly outward from near the lateral ocellus toalong the eye, on the frons the pattern radiates upward and outward from between theantennal sockets; ground shiny between wrinkles; middle with fine longitudinal wrinklesand incomplete fine frontal line; middle depressed from median ocellus to acme. Apex ofclypeus shallowly emarginate. Dorsal end of facial fovea extended to level of lateral ocellus.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct, fine pits 1-2 pitwidths apart, ground granularly coriaceous, dull; hair medium-length (8-12), white, mostlysimple. Scutellum slightly lustrous. Mesepisternum with distinct, round pits about 1 pitwidth apart, and short to medium-length, white, weakly plumose hair; ground roughenedto weakly striate, slightly lustrous. Hypoepimeron punctured, the pits partially obscuredby striate ground. Metanotum dull. Basal area of propodeum broadly convex, roughenedoverall by reticulate rugae except the posterior, vertical, coriaceous face; ground shinybetween rugae; lateral carina strong, terminating anterolaterally in a slightly protuberantarea.

Metasoma. Disk of T2 with distinct punctation; minute, shallow pits more than 2 pitwidths apart; smoothly coriaceous, shiny; with very short, fine, pale, appressed hair. S2without protuberance. T6 hair long, dark brown, erect.

Color. Body black. Face unmarked. Pronotum with collar broad, ivory transverse collarstripe complete; lobes unmarked. Legs blackish brown except: pale area on fore tibia; foreand mid tibiae marked full length with ivory stripe, hind tibia ivory except in caudal viewwith dark spot in apical half. Pale golden hairs dorsoapically on all tibiae. Wings darksmoky, some with reddish iridescence on the underside.

Perkins’ collections. Kauai: Makaweli, 2000 ft (BMNH, BPBM, OUMNH), >2000 ft(BPBM).

Recent collections. None.Flower records. None.Remarks. The most beautiful of the Hawaiian Hylaeus. Colors of the integument and

pubescence are based on old specimens.References. Perkins, 1910:604 (key); 1913:82 (biogeography).

Hylaeus (Nesoprosopis) psammobius (Perkins)(Fig. 56)

Nesoprosopis psammobia Perkins, 1911:724-725 (♂♀).Hylaeus (Nesoprosopis) psammobius, Meade-Waldo, 1923:35 (list).Nesoprosopis psammobia, Daly, 1994:118 (lectotype ♀ BPBM; Hawaii, Hilo coast;


DIAGNOSIS. Medium-sized bees with clear to smoky wings. Male face with yellow mark(clypeus, lower paraocular areas) below antennae partly divided by brown cuticle alongsulci, unique lower paraocular depression bordered along eye by a narrow, low ridge, andunique median process of S8 dilated at base, but very slender in distal half and with longtips. Female face with or without three separate small marks (clypeus with subapical yellow


stripe and yellow stripes along the eyes); legs with hind tibia with or without obscureyellow spot at very base.

Distribution. Hawaii and Maui (new record); coast. Very rare; no recent collectionsfrom Hawaii.

Male. Measurements: Malar space, 6; eye, 135:113:103:74; clypeus, 69×69; frons,22×25:87; interalveolar, 15:25:49:59, D 15; interocellar, 32:29:22:29, D 16; scape, 44×32;wing length, 4.3 mm (Hana, Maui, DOA).

Head. Malar space short (ratio about 0.4 DMO). Upper half of upper paraocular areaand frons uniformly with distinct, nearly round pits less than 1 pit width apart and withground smooth, lustrous; at middle pits smaller, contiguous, slightly distorted; frontal linefaint or absent. Lower half impunctate, area above antennal sockets shiny, extended dorsally

0.5 mm

A B C D

E F

1.0 mm

Fig. 56. Hylaeus psammobius, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


about half distance to median ocellus; lateral impunctate area with fine oblique ridges.Paraocular depression distinct, extended above antennal sockets and bordered along eyeby a narrow, low ridge. Frontoclypeal sulcus nearly straight. Scape in frontal view moderatelydilated, width more than half length, ratio about 0.7; lateral edge in frontal view straight;moderately arched in lateral view with maximum arch near the middle of the scape; undersideof median edge with a narrow line; with medium-length, pale brown, mostly simple hair.Vertex hair 12-20, brown, mixed simple and weakly plumose.

Mesosoma. Scutum with distinct pits 1 pit width apart or less, ground granular, dull toslightly lustrous; hair medium-length (8-16), pale brown, weakly plumose. Scutellum withpits slightly larger, ground slightly lustrous. Metanotum dull. Mesepisternum with distinct,round pits about 1 pit width apart, and long, yellowish white, plumose hair; ground minutelywrinkled, slightly lustrous. Hypoepimeron shallowly punctate, minutely granular, slightlylustrous, slightly swollen. Basal area of propodeum reticulately rugose or areolate to andincluding brow, granular between rugae.

Metasoma. Disk of T2 impunctate; minutely smoothly coriaceous, lustrous, metasomadistinctly hairy with fine, pale, moderately long, prostrate hairs spaced a hair length apart,longer and denser at sides, especially on T1. T7 hair short, brown, mostly prostrate.

Terminalia. Lateral process of S7 tapered, slender, longer than width of hemisternite.Median process of S8 weakly dilated or swollen at base and very narrow in distal half, apexbifurcate with long, rounded tips that are narrow at base and broad at apex. Apices ofgonoforceps somewhat pointed, longer than penis valves.

Color. Labrum and mandible dark brown. Facial marks appearing as 3 partly separateyellow marks, clypeus yellow except for narrow black margin along sulci at base and sides,lower paraocular areas with triangular marks almost reaching to antennal sockets,supraclypeal area with or without obscure yellow spot. Prothorax unmarked or with verysmall, narrow marks on lobes. Head and mesosoma black; metasoma blackish brown. Legsblack to dark brown, unmarked except fore tibia with pale area, mid and hind tibiae withobscure basal yellow dot. Wings slightly smoky.

Female. Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area andfrons with distinct, mostly round pits less than 1 pit width apart; ground smooth and shiny;at middle pits smaller and some contiguous, some weak longitudinal wrinkles at midline,with or without frontal line. Apex of clypeus shallowly emarginate. Dorsal end of facialfovea extended to level of lateral ocellus. Vertex hair 6-12, brown, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits about 1 pitwidth or less apart, ground slightly lustrous; hair medium (mostly even, about 6, a fewlonger to 12), pale yellowish brown, mixed weakly and densely plumose. Scutellum withslightly larger pits 1-2 pit widths apart, ground lustrous. Mesepisternum with indistinct,round pits, about 1 pit width or less apart, and long, yellowish white, weakly plumose hair;ground minutely coriaceous, slightly lustrous. Hypoepimeron obscurely punctured, slightlylustrous, not swollen. Metanotum slightly lustrous. Basal area of propodeum lineately rugoseor slightly reticulately rugose to brow, spaced 2-4 or more ruga widths apart, shiny betweenrugae; brow minutely wrinkled and slightly lustrous.


Metasoma. Disk of T2 impunctate, smoothly coriaceous, shiny; fine, pale, appressedhairs spaced about one hair length apart. T6 hair medium-length, brown, erect.

Color. Head and mesosoma dark brown to black (sometimes clypeus a lighter shade ofbrown than rest of head); metasoma dark brown. Face unmarked or with up to three smallmarks: clypeus with or without irregular, transverse, subapical yellow stripe; lowerparaocular area with or without irregular yellow stripe along the eye, not extended abovelevel of antennal socket; supraclypeal area and scape unmarked. Legs dark brown exceptfore tibia with pale area and obscure yellow spot at very base; hind tibia with or withoutobscure yellow spot at very base. Wings clear to slightly smoky.

Perkins’ collections. Hawaii: Hilo coast (BPBM, OUMNH).Other collections. Hawaii: Hilo coast, 3♀ VI-03 (Perkins, BPBM 3558); 1♂ 14-VI-?,

1♀ VIII-08 (W. M. Giffard, DOA). Maui: Hana, 1♂ 10-VI-1943 (N. L. H. Krauss, DOA).Recent collections. Maui: Eleilei Point, coast.Flower records. Bacopa monnieri, Sesuvium portulacastrum.Remarks. Windward coastal habitat is severely degraded, and it is possible that this

species may be extirpated on Hawaii. However, it may still survive in small patches ofnative vegetation. Other bees have not been observed visiting the plants it was found at onMaui, Bacopa and Sesuvium, and it may be necessary to look in different areas than thosewhere other coastal bees are typically found. The scutum pits are unusually close together,a character usually only found in the “punctured terga” species (but also in H. longiceps).

References. None.

Hylaeus (Nesoprosopis) pubescens (Perkins)(Fig. 57)

Nesoprosopis pubescens Perkins, 1899:107-108 (♂♀; Pl. 2, Fig. 21).Hylaeus (Nesoprosopis) pubescens, Meade-Waldo, 1923:34 (list).Nesoprosopis pubescens, Daly, 1994:118 (lectotype ♂ BMNH 17a 139a; Hawaii, Hilo,


DIAGNOSIS. Large bees with punctured terga, dark wings, face without marks; complete,incomplete, or no transverse stripe on the pronotal collar. Males with yellowish stripe onscape; and, especially in the female, a unique hump on second sternum seen in lateral view(see also remarks for sister species, H. fuscipennis).

Distribution. Hawaii; most often found in montane wet to mesic forest, occasionally indry shrubland.

Male. Measurements: Malar space, 7; eye, 179:146:143:101; clypeus, 100×90; frons,35×40:113; interalveolar, 20:36:59:72, D 28; interocellar, 33:41:38:49, D 19; scape, 61×49;wing length, 7.6 mm (Hawaii, N. Kona, BPBM 3548).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area with somewhatdistorted pits less than 1 pit width apart and separated by fine parallel wrinkles that curveacross the sides of the frons from the lateral ocellus to along the eye; frons with pitsrelatively distinct, nearly round, and with diminished wrinkles; ground generally slightly


lustrous, more lustrous near center; middle with some fine longitudinal wrinkles, with orwithout a fine frontal line; no impunctate area above antennal sockets. Lower paraoculardepression distinct, shallow to moderate. Frontoclypeal sulcus arched or nearly straight.Scape in frontal view moderately dilated, width more than half length, ratio about 0.8;lateral edge in frontal view distinctly curved (convex) or forming a broad obtuse angle inthe middle; strongly arched in lateral view with maximum arch near middle of scape;underside of median edge with a groove; with medium-length to long, yellowish white,mostly simple hair. Vertex hair 20-30, white, mostly simple.

Mesosoma. Scutum with distinct pits 1 pit width apart or less, ground minutely granular,dull to slightly lustrous; hair medium to long (8-14, few longer to 20), yellowish white,mixed simple and weakly plumose. Scutellum with pits slightly larger, lustrous.

0.5 mm

A B C D

E F

1.0 mm

Fig. 57. Hylaeus pubescens, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


Mesepisternum pits distinct, round, less than 1 pit width apart, and long, yellowish white,plumose hair; ground smooth to minutely wrinkled, lustrous. Hypoepimeron punctured,smooth, distinctly swollen. Metanotum dull. Basal area of propodeum slightly protuberantin middle, roughened overall by reticulate rugae except the posterior, vertical, coriaceousface, or roughened over entire surface; rugae and ground minutely wrinkled, slightly lustrous,some spaces between rugae shiny; lateral carina strong, terminating anterolaterally in aprotuberant area.

Metasoma. Disk of T2 with distinct punctation; fine pits 1-2 pit widths apart, and fine,pale, appressed hairs denser, longer, plumose, and white at posterolateral margins; groundsmoothly coriaceous, lustrous. T5-7 hair long, golden to orange-brown, prostrate.


Color. Labrum and mandible blackish brown to black. Face unmarked; scape in frontalview with yellowish stripe on lateral margin. Pronotum with ivory transverse collar stripevariable, complete to absent; lobes unmarked. Legs dark brown to black except: fore tibiawith pale area and with or without a basal ivory mark; all femora with or without obscureapical marks; mid tibia in frontal view with small basal ivory mark and sometimes withobscure apical ivory mark; mid basitarsus with basal ivory mark; hind tibia in frontal viewwith basal ivory mark up to half length of segment, and sometimes with obscure apicalivory mark; hind basitarsus with basal ivory mark up to full length of segment. Wings darksmoky, some bees with red and green iridescence along distal margins, reddish or reddishand bluish iridescence in center, especially the undersides.

Female. Head. Malar space medium (ratio about 0.6-0.7 DMO). Upper paraocular areawith shallow, slightly distorted pits about 1 pit width apart or less, some bees with more orless fine parallel wrinkles that curve across the sides of the area from near the lateralocellus to along the eye; frons with pits more distinct and round and with wrinkles lessevident; fine longitudinal wrinkles at middle, middle from median ocellus to acme notdepressed; with or without a fine frontal line. Apex of clypeus distinctly emarginate. Apexof mandible with three teeth. Dorsal end of facial fovea extended to level of lateral ocellus.Vertex hair 20-30, white, weakly plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground slightly granularly coriaceous, dull to slightly lustrous; hair white, mediumto long (10-20), weakly plumose. Scutellum with pits slightly larger, ground smoothlycoriaceous, lustrous. Mesepisternum with pits distinct, round, less than 1 pit width apart,and medium-length, yellowish white or brownish white, weakly plumose hair; ground smoothto minutely wrinkled, lustrous. Hypoepimeron punctured, smooth, swollen. Metanotumdull. Basal area of propodeum protuberant in middle, roughened overall by reticulate rugaeexcept the posterior, vertical, coriaceous face; rugae and ground minutely wrinkled, slightlylustrous, some spaces between rugae shiny; lateral carina strong, terminating anterolaterallyin a protuberant area.

Metasoma. Disk of T2 with distinct punctation; minute pits 1 or more pit widths apart,and fine, pale, appressed hairs becoming denser, longer, plumose, and brownish white at


posterolateral margins; ground smoothly coriaceous, lustrous. Sternum 2 with low, medianprotuberance. T5-6 hair long, golden to orange-brown, prostrate.

Color. Body and legs dark brown to black. Face and legs unmarked. Pronotum withnarrow transverse ivory collar stripe variable: nearly complete, interrupted, obscure, orabsent; lobes unmarked. Wings dark smoky, some bees with red and green iridescencealong distal margins, reddish iridescence medially, especially the undersides.

Perkins’ collections. Hawaii: Hilo, 2000 ft (BMNH, BPBM), 1800 ft (BPBM); Kau,4000 ft (BMNH, BPBM); Kilauea (BPBM), 4000 ft (OUMNH); Kona, N. (BPBM), 4000ft (BPBM); Kona, S., 2500 ft (BPBM); Olaa, 1500 ft (BPBM); Waimea, Kohala Mts.,4000 ft (BPBM); Puna, 2000 ft (BMNH).

Recent collections. Hawaii: Kau Desert Trail head, 930 m; Devastation Trail, 3700 ft;Hilina Pali Rd., 3300 ft; Mauna Loa Strip Road, 1200 m; Kipuka Ki, 4100 ft; Olaa ForestReserve, 915 m; Olaa Small Tract, 3800 ft; Puu Waawaa, 4100 ft; Kona Forest Unit ofHakalau National Wildlife Refuge camp, 5300 ft; Kau Forest Reserve, Mountain HouseRd., 3000 ft; Upper Hamakua Ditch, 3700 ft.

Flower records. Cheirodendron, Hypochoeris radicata*, Metrosideros, Myoporum,Psychotria hawaiiensis, Santalum paniculatum, Tibouchina urvilleana*.

Remarks. This is the most abundant of the large species, commonly found throughoutthe island of Hawaii in wet and mesic forest. It is strongly associated with Metrosideros.The dense hair at the apex of the metasoma is conspicuous and distinctive.

References. Perkins, 1907b: 97 (record: Hawaii, Kilauea); 1910:604 (key). Bridwell,1919:135 (morphology). Timberlake, 1919a: 225 (parasitoid: Nesencyrtus kaalae; record:Hawaii, Kilauea); 1919b: 185 (parasitoid: Nesencyrtus kaalae; record: Hawaii, Kilauea).Cockerell, 1926:308 (record: Hawaii, Volcano House). Williams, 1927:428-429 (nest: figure,in decayed Metrosideros polymorpha; parasitoid: Nesencyrtus kaalae; record: Hawaii,Kilauea, 4000 ft). Swezey, 1954:125 (nest: beetle burrows in Metrosideros collinapolymorpha). Beardsley, 1976:188, 209 (parasitoid: Coelopencyrtus kaalae). Daly andCoville, 1982:75-81 (description of larva; trap nests; nest: in beetle burrows inLeptospermum scoparium and Acacia koa; flower: Hypochoeris radicata; parasitoid:Coelopencyrtus kaalae and C. sexramosus; inquiline: Afrocypholaelaps africana; record:Hawaii, Kilauea). Suehiro, 1986:61 (record: Hawaii, Kona).

Hylaeus (Nesoprosopis) rugulosus (Perkins)(Fig. 58)

Nesoprosopis rugulosa Perkins, 1899:84 (♀).Hylaeus (Nesoprosopis) rugulosus, Meade-Waldo, 1923:34 (list).Prosopis ovatula Warncke, 1970:748 (unwarranted replacement name for Nesoprosopis

rugulosa Perkins) (new synonymy)Nesoprosopis rugulosa, Daly, 1994:118 (lectotype ♀ BMNH 17a 2750; Hawaii, Kau,


DIAGNOSIS. Medium-sized bees with smoky wings and basal area of propodeumreticulately rugose to brow. Male black and unmarked except pale area on fore tibia;


supraclypeal area much wider than long, large shiny area above antennal sockets; scapemoderately dilated even at base; gonoforceps uniquely truncate at apex. Female black andunmarked; with unique mandible straight across at apex, as wide or wider at apex than atarticulation.

Distribution. Hawaii; montane wet forest.Male. Measurements: Malar space, 5; eye, 140:123:123:78; clypeus, 66×69; frons,

22×35:91; interalveolar, 17:27:54:64, D 17; interocellar, 32:33:29:32, D 16; scape, 44×29;wing length, 4.5 mm (Hawaii, Kipuka Nene, 2900 ft).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area and frons withround pits less than 1 pit width apart, even near the middle, but at the very middle becomingsmaller, contiguous, and distorted, and obscuring part of frontal line; ground minutelywrinkled and slightly lustrous between pits. Large impunctate, shiny area above antennalsocket. Lower paraocular depression distinct, especially evident along subantennal sulci.Frontoclypeal sulcus slightly arched. Supraclypeal area much wider than long; W/L ratioabout 1.7. Scape in frontal view moderately dilated and wide even near base, width morethan half length, ratio about 0.7; lateral edge in frontal view straight; moderately arched inlateral view with maximum arch near middle of scape; no groove on underside; with long,yellowish white, weakly plumose hair. Vertex hair 16-24, brownish white, mixed simpleand plumose.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground lustrous; hair medium-long(10-20), pale brown, weakly plumose. Scutellum with slightly larger pits, ground lustrous.Mesepisternum with indistinct, shallow, round pits, about 1 pit width apart, and long,white, plumose hair; ground minutely coriaceous, lustrous. Hypoepimeron impunctate,lustrous, slightly swollen. Metanotum dull. Basal area of propodeum reticulately rugose tobrow; shiny between rugae; brow wrinkled coriaceous and lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, palebrownish, appressed hairs spaced about one hair length apart or less. T7 hair short, brown,prostrate.

Terminalia. Lateral process of S7 tapered, narrow, and with length slightly longer thanwidth of hemisternite. Median process of S8 moderately dilated, apex bifurcate with narrow,rounded tips. Apices of gonoforceps truncate, subequal to penis valves.

Color. Head and mesosoma black; metasoma brownish black. Labrum and mandibleblack. Face and scape unmarked; antenna yellow to brown beneath. Pronotum unmarked.Legs black; unmarked except fore tibia with pale area and fore basitarsi dull orange yellow.Wings smoky and slightly iridescent.

Female. Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular areaand frons rather uniformly punctured with round pits less than 1 pit width apart; groundminutely wrinkled, slightly lustrous; middle slightly depressed, with faint frontal line. Apexof clypeus nearly straight. Dorsal end of facial fovea extended to level of lateral ocellus.Mandible straight across at apex, widest at apex (Fig. 9C). Vertex hair 16-24, brownishwhite, mixed simple and plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with minute, shallow pits 2 ormore pit widths apart, ground slightly, granularly coriaceous, slightly lustrous; hair medium


to long (8-20), pale brown, simple. Scutellum with pits slightly larger. Mesepisternum withindistinct, shallow, nearly round pits 1 pit width apart, and medium-length, weakly plumose,white hair; ground nearly smoothly coriaceous, slightly lustrous. Hypoepimeron obscurelypunctured, ground smoothly coriaceous, lustrous, nearly flat. Metanotum dull. Basal areaof propodeum rather swollen, convex, with short anterior area, reticulately rugose to browwith some long, transverse rugae; weakly rugose beyond brow.

Metasoma. Disk of T2 impunctate, lustrous, with very short, appressed hairs spacedmore than a hair length apart. T6 hair medium-length, black, erect.

Color. Head and mesosoma black; legs dark brown; unmarked. Wings clear to smoky.Perkins’ collections. Hawaii: Kau, 4000 ft (BMNH); Kilauea (BPBM).

0.5 mm

A B C D

E F

1.0 mm

Fig. 58. Hylaeus rugulosus, new combination, male. A. Seventh sternum, ventral view. B. Eighthsternum, ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsalview. E. Face with left scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extentof pubescence shown by dashed lines in B-E.


Recent collections. Hawaii: Napau Trail, 2800 ft; Kipuka Nene, 2900 ft; Puu Puai,3700 ft; below 3-Tree Kipuka, 1890 m.

Flower records. Metrosideros.Remarks. This species has been rarely collected, but a large series was recently collected

around a stone structure, apparently a nesting site. As pointed out by Snelling (1975), theNearctic species often known as Hylaeus rugulosus (now regarded as a synonym of H.modestus citrinifrons) is correctly spelled H. rugosulus (Cockerell, 1896).

References. Perkins, 1907b: 97 (record: Hawaii, Kilauea). Fullaway, 1918:393-394 (♂).

Hylaeus (Nesoprosopis) satelles (Blackburn)(Fig. 59)

Prosopis satellus Blackburn, 1886:140-141, 148 (♂; holotype ♂ BPBM 2340, Maui,Haleakala, about 5000 ft; wing length, 6.5 mm, tip damaged. Blackburn used both P.satellus and P. satelles; Perkins, 1899, as first reviser, used P. satelles).

Nesoprosopis satelles, Perkins, 1899:109-110 (♂♀; Pl. 2, Fig. 23).Hylaeus (Nesoprosopis) satelles, Meade-Waldo, 1923:35 (list).Nesoprosopis satelles, Daly, 1994:114 (holotype ♂ confirmed).

DIAGNOSIS. Large bees with punctured terga, dark wings, and propodeum sparselysculptured. Male with scape extremely dilated, circular. Similar to H. muranus; see remarksfor that species.

Distribution. Maui, Lanai, and Molokai; montane wet forest. No recent collections.Male. Measurements. Malar space, 8; eye, 154:146:146:97; clypeus, 92×81; frons,

23×33:97; interalveolar, 26:33:59:67, D 26; interocellar, 33:41:26:33, D 18; scape, 54×61;wing length, 6.3 mm (Lanai, Halepaakai, 3000 ft; BPBM 3560).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area and frons withshallow, slightly distorted pits less than 1 pit width apart, pits about the same size andspacing from the sides to near the middle, ground minutely wrinkled, sometimes shiny,generally lustrous; at middle a fine frontal line that may be partly obscured by adjacentpunctation. Lower paraocular depression distinct, extended above antennal socket, andshiny next to eye. Frontoclypeal sulcus slightly arched. Scape extremely dilated, widerthan long, ratio about 1.1; nearly circular in frontal view; frontal surface unevenly convexand uniformly covered with fine pits less than 1 pit width apart; strongly arched in lateralview with maximum arch near middle of scape; with medium-length, yellowish brown,simple hair. Vertex hair medium-length, yellowish brown, mostly simple.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground lustrous; hair medium-length(8-12), yellowish white, mostly simple. Mesepisternum with distinct, round pits about 1 pitwidth apart, and medium-length, pale yellowish brown, simple and weakly plumose hair;ground smooth, lustrous. Hypoepimeron indistinctly punctured, smooth, lustrous, swollen.Metanotum punctation obscure, dull. Basal area of propodeum at the center of the frontwith sparse, short, lineate rugae and at the sides of the front, a few longer, oblique, lineaterugae (rugae stronger in Molokai specimen); brow smoothly coriaceous and lustrous.


Metasoma. Disk of T2 with minute, shallow pits about 1 pit width apart, smoothlycoriaceous and lustrous; fine, pale, appressed hair. T7 hair long, brown, erect, simple.

Terminalia. Lateral process of S7 tapered, length subequal to width of hemisternite.Median process of S8 strongly dilated, dilation membranous, apex bifurcate with broadtips. Apices of gonoforceps subequal to penis valves.

Color. Head and mesonotum dark brown to black; pleura, propodeum, and metasomareddish brown to dark reddish brown. Labrum and mandible dark reddish brown to black.Face marks variable: clypeus entirely reddish yellow or only at apex; paraocular areasunmarked; and supraclypeal area reddish yellow to acme or obscurely so. Scape variable,dark or with the lateral margin reddish yellow. Pronotum unmarked. Legs reddish brownto dark brown except fore femur, tibia, and basitarsus with pale areas. Wings dark brown.

Fig. 59. Hylaeus satelles, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.

E

0.5 mm

A B C D

F


Female. Head. Malar space medium-length (ratio about 0.5 DMO). Upper paraoculararea and frons with shallow, slightly distorted pits less than 1 pit width apart, pits about thesame size and spacing from the sides to near the middle, ground minutely wrinkled orsometimes shiny, generally lustrous; middle with a fine frontal line that may be partly obscuredby adjacent punctation. Apex of clypeus very shallowly emarginate. Dorsal end of facialfovea extended to level of lateral ocellus.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground lustrous; hair medium to long (8-16), yellowish white, mostly simple.Mesepisternum with distinct, round pits about 1 pit width apart, and brownish white, plumosehair; ground smooth, lustrous. Hypoepimeron distinctly punctured, smooth, lustrous,swollen. Metanotum slightly lustrous. Basal area of propodeum nearly glabrous or withweak, lineate rugae less than halfway to brow; brow smooth coriaceous, lustrous.

Metasoma. Disk of T2 with minute, shallow pits about 1 pit width apart, smoothlycoriaceous and lustrous; fine, pale, appressed hair, slightly denser along apical margin. T6hair mostly simple except minutely plumose at sides, long, black, erect.

Color. Body dark reddish brown to black. Face marks: clypeus and (in some specimens)adjacent supraclypeal area reddish yellow. Scape almost entirely reddish yellow, or markedonly along lateral edge. Legs dark brown except: fore femur and tibia with pale areas.Wings dark brown.

Perkins’ collections. Maui: Haleakala, 5000 ft (BMNH, BPBM). Lanai: center ofLanai, forests above Koele, 2000 ft (OUMNH); Halepaakai, 3000 ft (BMNH, BPBM);“Lanai,” 2000 ft (BMNH, BPBM, OUMNH); Mts. Koele, 2000 ft (BMNH, BPBM), >2000ft (BPBM). Molokai: “Molokai,” 4000 ft (OUMNH); Molokai Mts., 4000 ft (BPBM),>4000 ft (BMNH, BPBM), east, 4000 ft (OUMNH).

Recent collections. None.Flower records. None.Remarks. Colors of the integument and pubescence are based on old specimens. Perkins’

specimens from Molokai are darker (entire mesosoma dark brown) than those from Lanai.References. Blackburn, 1887:200-201, 208 (♂, key). Dalla Torre, 1896:32 (list). Perkins,

1910:604 (key); 1913:79 (ecology). Fullaway, 1918:396 (record: Maui, Iao Valley).

Hylaeus (Nesoprosopis) setosifrons (Perkins)(Fig. 60)

Nesoprosopis setosifrons Perkins, 1899:108-109 (♂♀; Pl. 2, Fig. 22; flower:Metrosideros, Rubus, Santalum).

Hylaeus (Nesoprosopis) setosifrons, Meade-Waldo, 1923:34 (list).Nesoprosopis setosifrons, Daly, 1994:119 (lectotype ♂ BMNH 17a 142a; Hawaii, Kau,


DIAGNOSIS. Large bees with punctured terga and dark wings. Male scape moderatelydilated, face with ivory marks in lower paraocular areas and sometimes subapically onclypeus. Female pronotal collar with transverse ivory stripe complete or interrupted; tibiae


with ivory stripes; basitarsi with or without ivory stripes. Both sexes with unusual dark,erect setae on face, almost all body hair black.

Distribution. Hawaii; montane mesic and wet forest.Male. Measurements: Malar space, 10; eye, 148:141:136:97; clypeus, 87×82; frons,

26×35:100; interalveolar, 23:31:54:64, D 26; interocellar, 29:41:18:31, D 17; scape, 49×39;wing length, 6.0 mm (Hawaii, Hilo, 2000 ft, BPBM 3574).

Head. Malar space medium-length (ratio about 0.6 DMO). Upper paraocular area withshallow, somewhat indistinct, round pits 1 pit width or less apart; frons with pits smallermedially, closer, shallower, but mostly round; ground minutely, irregularly wrinkled amongpits and across the middle, dull; very small impunctate area above antennal socket. Lowerparaocular depression distinct, but shallow. Frontoclypeal sulcus slightly arched. Face withunusual dark, erect setae. Scape in frontal view moderately dilated, width more than halflength, ratio about 0.8; lateral edge in frontal view slightly convex; moderately arched inlateral view with maximum arch near middle of scape; no groove on underside; with long,brown, mostly plumose hair. Vertex hair 20-30, black, mostly simple.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground granularly coriaceous, slightlylustrous; hair long (12-20 or more, somewhat shorter in small specimens), black, mostlysimple. Scutellum with slightly larger pits, smoothly coriaceous, lustrous. Mesepisternumwith indistinct, shallow pits about 1 pit width apart, and long, brown, weakly plumose hair;ground nearly smooth, slightly lustrous. Hypoepimeron punctured, nearly smooth, slightlylustrous. Metanotum dull. Basal area of propodeum convex in middle but not distinctlyprotuberant; irregularly roughened or lineately rugose almost to brow, rugae spaced 2-4rugae apart; shiny between rugae; ground of brow and below varying from smoothlycoriaceous and lustrous to wrinkled and dull. Lateral carina strong; without an anterolateralprotuberance.

Metasoma. Disk of T2 with punctation of indistinct, shallow pits more than 2 pit widthsapart; smoothly coriaceous, lustrous; with very short, fine, brown, appressed hair. T7 hairlong, black, prostrate to erect.

Terminalia. Lateral process of S7 tapered, with length subequal to width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with broad, rounded tips. Apicesof gonoforceps slightly longer than penis valves.

Color. Labrum and mandible blackish brown to black. Face with two or three marks:clypeus unmarked or with an apical transverse stripe; lower paraocular area largely orentirely ivory from just below antennal socket; supraclypeal area and scape unmarked.Pronotum unmarked. Legs dark brown to black except: fore tibia with pale area combinedwith irregular dorsal yellowish stripe; mid tibia with a basal ivory or yellowish trace,basitarsus with basal ivory or yellowish marking; hind tibia with irregular ivory stripe inbasal half, often extended to apex dorsally. Pubescence generally dark brown to black,except on propodeum and legs. Wings dark smoky with faint red and green iridescencealong distal margins.

Female. Head. Malar space medium-length (ratio about 0.6 DMO). Upper paraoculararea and frons with shallow, slightly distorted pits, less than 1 pit width apart and more orless aligned in rows separated by fine, parallel wrinkles (wrinkles stronger or weaker in


0.5 mm

A B C D

E F

1.0 mm

Fig. 60. Hylaeus setosifrons, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.

different bees); at the sides the wrinkles curve broadly outward from near the lateral ocellusto along the eye, whereas on the frons the pattern radiates upward and outward frombetween the antennal sockets; middle with fine longitudinal wrinkles with or without a finefrontal line; middle from median ocellus to acme slightly depressed. Apex of clypeusshallowly emarginate. Face with unusual dark, erect setae. Dorsal end of facial foveaextended to level of lateral ocellus. Vertex hair 20-30, black, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground granularly coriaceous, dull; hair long (14-20 or more), black, mostly simple.Scutellum with slightly larger pits, smoothly coriaceous ground, slightly lustrous.Mesepisternum variable with distinct or indistinct, shallow pits about 1 pit width apart, andbrownish white, plumose hair; ground variable, nearly smooth, minutely wrinkled to weakly


striate, slightly lustrous. Metanotum dull. Basal area of propodeum broadly convex,roughened overall by irregularly lineate or reticulate rugae except the posterior, vertical,coriaceous face; rugae and ground minutely wrinkled, dull or slightly lustrous; lateral carinastrong, terminating anterolaterally in a slightly protuberant area. Hypoepimeron punctured,the pits partially obscured by minute, parallel, longitudinal wrinkles.

Metasoma. Disk of T2 with distinct punctation; minute, shallow pits more than 2 pitwidths apart; smoothly coriaceous, shiny; with very short, fine, light brown, appressedhair. T6 hair long, black, erect.

Color. Head and mesosoma black; metasoma dark brown to black. Face unmarked.Pronotum with ivory transverse collar stripe complete or interrupted; lobes unmarked.Legs dark brown except: pale area on fore tibia plus ivory stripe nearly full length ofsegment; fore basitarsus with or without ivory at base; mid and hind tibiae and hind basitarsuswith ivory stripe nearly full length of segments; mid basitarsus at base with or withoutivory marks. Wings dark smoky, some with red or bluish iridescence.

Perkins’ collections. Hawaii: Hilo, 2000 ft (BPBM); Kau, 4000 ft (BMNH, BPBM);Kilauea (BPBM), 4000 ft (OUMNH); Kona, 2000 ft, 3000 ft, 4000 ft (BPBM); Kona, N.(OUMNH), 3000 ft (OUMNH).

Recent collections. Hawaii: Mauna Loa Estates, 3600 ft; Tree Molds, 4000 ft;Earthquake Trail, 4000 ft; Volcano, Old Japanese Schoolhouse, 3750 ft.

Flower records. Cheirodendron, Metrosideros, Psychotria hawaiiensis, Rubus sp.,Santalum paniculatum.

Remarks. This species is widely distributed in the wet and mesic forest around Kilauea,but is uncommon. Nearly all of the body hair is black, and the erect clypeal hair is distinctive.

References. Perkins, 1907b: 97 (record); 1910:604 (key); 1913:81 (parasitoid:Eupelmus). Fullaway, 1918:398 (comparison with N. anomala). Suehiro, 1986:61 (record:Hawaii, Kona).

Hylaeus (Nesoprosopis) simplex (Perkins)(Fig. 61)

Nesoprosopis simplex Perkins, 1899:79 (♂♀; Pl. 2, Fig. 9).Hylaeus (Nesoprosopis) simplex, Meade-Waldo, 1923:33 (list).Nesoprosopis simplex, Daly, 1994:119 (lectotype ♂ BMNH 17a 157b; Hawaii, Kau,


DIAGNOSIS. Medium-sized bees with smoky wings. Male face with oval yellow markdivided by fine black sulci, legs unmarked, unusual lateral process of S7 long and broad,median process of S8 not dilated and with narrow tips, unusual gonoforceps expandedapically beyond the penis valves and with pubescence extended around apex to inner angle.Female black and unmarked. Closely related to H. chlorostictus and H. facilis.

Distribution. Hawaii; lowland and montane dry shrubland and mesic forest. No recentcollections.


Male. Measurements: Malar space, 7; eye, 143:123:115:79; clypeus, 69×69; frons,29×28:100; interalveolar, 17:27:54:64, D 22; interocellar, 32:31:23:28, D 17; scape, 51×27;wing length, 4.8 mm (Hawaii, Puna, 1000 ft, BPBM 3593).

Head. Malar space short (ratio about 0.4 DMO). Upper paraocular area and frons ratherflat and with indistinct, mostly round, shallow pits 1-2 pit widths apart even at the middle,ground minutely wrinkled, the wrinkles more or less parallel transversely across the middleand longitudinally at sides along eye, slightly lustrous; median, impunctate, slightly depressed,lustrous area above antennal sockets. Lower paraocular depression distinct, but shallow.Frontoclypeal sulcus straight to slightly arched. Scape in frontal view slightly dilated, widthhalf length, ratio about 0.5; lateral edge in frontal view nearly straight; strongly arched inlateral view with maximum arch near middle or slightly toward apex of scape; no grooveon underside; with medium-length, pale yellowish brown, weakly plumose hair. Vertex hair16-30, brown, plumose.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground slightly granular and slightlylustrous; hair medium-length (12-20), pale yellowish brown, plumose. Scutellum with slightlylarger pits, smoothly coriaceous, lustrous. Mesepisternum with indistinct, shallow pits about1 pit width apart, and long, yellowish white, weakly plumose hair; ground minutelycoriaceous to striate, slightly lustrous. Hypoepimeron impunctate, nearly smooth, lustrous,slightly swollen. Metanotum dull. Basal area of propodeum lineately rugose to brow, shinybetween rugae, weakly reticulate at brow; brow below smoothly coriaceous, slightly lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with short, fine, pale,appressed hairs spaced more than a hair length apart. T7 hair medium-length, brown,prostrate.

Terminalia. Lateral process of S7 broad, sword-shaped, length much greater than widthof hemisternite. Median process of S8 linear, apex bifurcate with long, filamentous, denselyhairy tips. Apices of gonoforceps exceeding penis valves.

Color. Labrum and mandible yellowish brown to brown. Face with a single large spotnot extended at the sides above the antennal sockets and with fine black sulci in “H” shape:clypeus entirely yellow or nearly so; lower paraocular area yellow below antennal socketand along sulci; supraclypeal area to acme yellow; scape unmarked. Pronotum unmarked.Legs dark brown except: fore tibia with pale area, with or without obscure basal yellowspots or stripes on all tibiae and fore basitarsus. Wings smoky.

Female. Head. Malar space medium-length (ratio about 0.5 DMO). Upper paraoculararea and frons with indistinct, mostly round pits about 1 pit width apart; ground minutelywrinkled, slightly lustrous; fine frontal line in some bees. Apex of clypeus shallowlyemarginate. Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 14-24, brown, plumose.

Mesosoma. Fore tarsus with hair apically curved. Scutum with pits 1-2 pit widths apart,ground granularly coriaceous, slightly lustrous; hair medium-length (10-14) yellowish white,plumose. Scutellum with pits slightly larger, smoothly coriaceous, lustrous. Mesepisternumwith indistinct pits about 1 pit width apart, and long, yellowish white, weakly plumosehair; ground minutely coriaceous to striate, slightly lustrous. Hypoepimeron impunctate,


nearly smooth, lustrous, slightly swollen. Metanotum dull. Basal area of propodeum lineatelyrugose to brow; shiny between rugae; brow granular to smoothly coriaceous, dull to slightlylustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, with fine, short, pale hairmore than a hair length apart. T6 hair medium-length, brown, prostrate.

Color. Head and mesosoma black; legs and metasoma dark brown; unmarked. Wingsdark smoky.

Perkins’ collections. Hawaii: Hamakua, 2000 ft (BPBM); Hilo (BMNH, BPBM), 4000ft (BPBM); Kau (BPBM), 4000 ft (BMNH, BPBM); Kilauea (BPBM), 4000 ft (OUMNH);Kona (BPBM), 4000 ft (BMNH, BPBM); Kona, N., 3000 ft (BPBM); Puna (BPBM),2000 ft (BMNH, BPBM, OUMNH).

0.5 mm

A B C D

E F

1.0 mm

Fig. 61. Hylaeus simplex, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.


Recent collections. None.Flower records. None.Remarks. Colors of the integument and pubescence are based on old specimens. See

remarks for H. facilis.References. Perkins, 1907b: 97 (record: Hawaii, Kilauea); 1910:601 (key). Fullaway,

1918:393 (record: Hawaii, Hilea, Kau, Honuapo, Kilauea-Honuapo auto road). Suehiro,1986:41, 61 (record: Hawaii, Kona).

Hylaeus (Nesoprosopis) solaris Magnacca and Daly, new species(Fig. 62)

DIAGNOSIS. Medium-sized bees with clear wings. Male with the face entirely yellow,broadly above antennal sockets, with or without a mark on the dilated scape; supraclypealarea longer than wide. Female black, unmarked; differing from the sympatric H. flavifronsand H. connectens by the short, golden hair of T6, and from H. chlorostictus by the veryshort pubescence of the scutum.

Distribution. Kauai; coast.Male. Measurements: Malar space, 11; eye, 148:100:92:71; clypeus, 78×68; frons,


Head. Malar space long (ratio about 0.8 DMO). Upper paraocular area and frons withdistinct, round pits, up to 1 pit width apart, smaller and closer in middle but not contiguous;ground smoothly coriaceous and lustrous over entire area; large impunctate area aboveantennal socket. Lower paraocular depression very shallow, indistinct. Frontoclypeal sulcusnearly straight; supraclypeal area much longer than wide, W/L ratio 0.6. Scape in frontalview moderately dilated, W/L ratio 0.7-0.8, greatest width just past middle; lateral edge infrontal view straight or very slightly concave; moderately arched in lateral view withmaximum arch around middle; underside of median edge with a long, narrow line; long,yellowish white, simple hair along median edge. Vertex hair 6-10, white to pale brown,plumose.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground smoothly coriaceous, slightlylustrous; hair medium-length (8-12), white, plumose. Scutellum with slightly larger pits,slightly lustrous. Mesepisternum with round, shallow pits about 1 pit width apart, groundsmoothly coriaceous and slightly lustrous, and with long, brownish white, plumose hair.Metanotum roughened, slightly lustrous. Basal area of propodeum irregularly, lineatelyrugose at front, shining between rugae, reticulate behind, brow minutely granular, lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart or more. T6 hair long, yellowish togolden, prostrate.

Terminalia. Lateral process of S7 tapered, broadly triangular, length subequal to widthof hemisternite. Median process of S8 long, strongly arched, evenly but weakly dilated;apex bifurcate, tips long, bent dorsad. Apices of gonoforceps subequal to penis valves.


Color. Labrum and mandible dark brown to black. Face with a single large mark: clypeusentirely yellow, with paraocular marks extended above antennal sockets in a broad line;supraclypeal area yellow. Scape with or without a broad stripe or spot on lateral half.Pronotum unmarked. Legs black except for pale area of front tibia, with or without smallspots at bases of all tibiae. Wings clear.

Female. Head. Malar space long (ratio about 0.8 DMO). Upper paraocular area andfrons with distinct, round pits, up to 1 pit width apart, evenly spaced, smaller and closeronly at middle; ground smoothly coriaceous and lustrous over entire area; impunctate areaabove antennal socket small. Upper portion of frontal line faint, forming a distinct grooveup to halfway between acme and median ocellus. Apex of clypeus slightly emarginate.Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 4-8(-12), yellowishwhite, plumose.

Fig. 62. Hylaeus solaris, new species, male. A. Seventh sternum, ventral view. B. Eighth sternum,ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E.Face with left scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E.

A B C D

0.5 mm

1.0 mm

E F


Mesosoma. Fore tarsus with hair apically curved. Scutum with distinct pits 1-2 pit widthsapart, ground slightly lustrous; hair short (6-8), yellowish white, plumose. Scutellum withslightly larger pits, slightly lustrous. Metanotum granular, dull. Mesepisternum with shallowpits 1-2 pit widths apart, ground smoothly coriaceous, lustrous; with short to medium,yellowish white, plumose hair. Hypoepimeron weakly punctate, striate, nearly flat; weaklyseparated from mesepisternum. Basal area of propodeum irregularly, lineately rugose atfront, shining between rugae, reticulate behind, brow minutely granular, lustrous.

Metasoma. Disk of T2 impunctate, almost smooth, shiny, with fine, pale, appressedhairs spaced about one hair length apart or more. T6 hair short, golden, prostrate.

Color. Head, mesosoma, metasoma, and legs black, except antennal flagellum light brownto bright yellow underneath, front tibia with small pale area.

Holotype ♂ labeled “Kauai, Polihale State Park, coastal dunes, on Scaevola sericea. K.Magnacca, 25 Aug 1999, KM-99-234” (CUIC).

Allotype ♀ with identical data (CUIC).Paratypes. Kauai, Polihale State Park, coast, 3♂ 4♀ 25-VIII-1999, on Scaevola sericea

(KM-99-234) (CUIC); 2♂ 2♀ with identical data (BPBM); Keoneloa Bay, coast just inlandeast of Hyatt Regency Hotel, 3♂ 2 ♀ 19-VIII-1989, J. A. Powell (UCB).

Flower records. Scaevola sericea, Sida fallax.Etymology. The species epithet solaris refers to the intense sun of the coastal habitat in

which the bees were collected, and to the bright yellow facial markings.Remarks. The relative abundance of this species at easily accessible locations makes it

surprising that it has not been collected before. However, its numbers at Polihale (as is thecase with other Hylaeus there) appear to be extremely seasonal. Like other coastal species,it is restricted in range; it is currently known only from Polihale and Keoneloa Bay. Hylaeussolaris shares with H. inquilina short vertex hair, broad paraocular stripes, and weaklyreticulate propodeum.

Hylaeus (Nesoprosopis) specularis (Perkins)(Fig. 63)

Nesoprosopis specularis Perkins, 1899:93-94 (♀; holotype ♀ BMNH 17a 2751; Hawaii,Kilauea; wing length, 3.8 mm).

Hylaeus (Nesoprosopis) specularis, Meade-Waldo, 1923:34 (list).Nesoprosopis specularis, Daly, 1994:119 (holotype ♀ confirmed).Nesoprosopis homoeochroma Perkins, 1911:722-723 (♂; type not located; Hawaii,

Kilauea) (new synonymy).Hylaeus (Nesoprosopis) homoeochromus, Meade-Waldo, 1923:35 (list).Nesoprosopis homoeochroma, Daly, 1994:116 (type not located).

DIAGNOSIS. Small bees with slightly smoky wings. Male face with three nearly separatemarks (clypeus except the basal and lateral edges and paraocular areas up to or slightlyabove antennal sockets, but not the supraclypeal area); frons with dark spot in middle;scape moderately dilated and with long hair along median edge; pronotum and legs marked


with yellow. Closely related to H. crabronoides, but without a thick head and with a largeimpunctate area above antennal socket. Female face with lower paraocular area with irregulartriangular yellow mark along the eye and up to level of antennal socket, rarely with atransverse apical mark on the clypeus; pronotum with yellow stripes on collar at sides andyellow lobes; legs with small, basal yellow spots on tibiae. Similar to H. dumetorum, butwith the upper frons smooth and lustrous; see remarks for that species.

Distribution. Hawaii, Oahu (new record), and Kauai (new record), possibly Maui;montane wet to mesic forest.

Male. Measurements: Malar space, 3; eye, 90:90:83:52; clypeus, 47×46; frons, 15×17:70;interalveolar, 15:18:41:49, D 15; interocellar, 24:26:13:23, D 13; scape, 32×23; wing length,3.6 mm, estimated (Hawaii, Kilauea, 4000 ft, BPBM 3602).

Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular area at thesides with nearly round pits less than 1 pit width apart; frons with pits becoming crowdedmedially and distorted in subpolygonal shapes; a dark rhomboid spot at the middle createdby dense black or dark brown hairs and very dense punctation; large impunctate shiny areajust below spot and above antennal socket. Lower paraocular depression shallow, sometimesindistinct. Frontoclypeal sulcus nearly straight. Scape in frontal view moderately dilated,width more than half length, ratio about 0.7; lateral edge in frontal view nearly straight;slightly arched in lateral view with maximum arch near middle of scape; underside of medianedge with a groove; with long, simple, pale brownish hair. Vertex hair 10-22, pale brown,simple.

Mesosoma. Scutum with indistinct, shallow pits about 2 pit widths apart, ground granular,dull; hair medium-length (8-14), pale brown, simple. Scutellum pits slightly larger, groundsmoothly coriaceous and slightly lustrous. Mesepisternum with indistinct to barely discerniblepits about 1-2 pit widths apart, and long, yellowish white, weakly plumose hair; groundminutely coriaceous, lustrous. Hypoepimeron impunctate, smooth, lustrous, slightly swollen.Metanotum slightly lustrous. Basal area of propodeum with rather long anterior area; sparse,weak, lineate rugae at front and extended about a quarter or less to brow, brow smoothlycoriaceous and lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, shiny, with fine, pale, appressedhairs spaced about one hair length apart. T7 hair long, dark brown, erect.

Terminalia. Lateral process of S7 tapered, with length subequal to width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with short, rounded tips. Apicesof gonoforceps longer than penis valves, curved inward.

Color. Labrum and mandible blackish brown to black. Face with three marks: clypeuslargely yellow except dark margin basally along epistomal sulcus above tentorial pits;paraocular area largely yellow to level of antennal socket or slightly above; supraclypealarea and scape unmarked. Pronotum with transverse collar stripe interrupted medially;lobes yellow. Fore leg dark brown except femur with apex yellow; tibia with pale area anddorsal yellow stripe; basitarsus mostly yellow; terminal tarsal segments yellowish brown tolight brown. Mid leg dark brown to black except: femur with very apex yellow; tibia withbasal and sometimes distal yellow spots; basitarsus mostly yellow. Hind leg dark brown toblack except: tibia with yellow stripe in basal third; basitarsus yellow in basal half. Some


Kauai specimens lack marks on lobes and have leg markings reduced to obscure spots.Wings slightly smoky.

Female. Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area andfrons convex; upper paraocular area with nearly round pits less than 1 pit width apart,ground lustrous to shiny; frons with pits crowded medially and partly distorted in shape,ground distinctly duller between antennal sockets and median ocellus, and at the middle afew fine, longitudinal carinulae; frontal line, if any, distinct only near acme; small shiny areajust above antennal socket. Apex of clypeus shallowly emarginate. Dorsal end of facialfovea extended to level of lateral ocellus. Vertex hair 6-12(-20), pale brown, simple.

Mesosoma. Fore tarsus with hair apically curved. Scutum with indistinct, shallow pitsabout 2 pit widths apart, ground granular, dull to slightly lustrous; hair short (4-10), brown,

0.5 mm

A B C D

E F

1.0 mm

Fig. 63. Hylaeus specularis, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


simple. Scutellum pits slightly larger, ground smoothly coriaceous and slightly lustrous.Mesepisternum with barely discernible pits about 2-3 pit widths apart, and long, yellowishwhite, weakly plumose hair; ground minutely coriaceous, lustrous. Hypoepimeronimpunctate, smooth, lustrous, slightly swollen. Metanotum dull. Basal area of propodeumwith sparse, weak, lineate rugae at front and extended about a quarter or less to brow;brow smoothly coriaceous and lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart. T6 hair long, dark brown, erect.

Color. Head and mesosoma dark brown to black; metasoma dark brown to black. Facemarks: clypeus with or without thin, apical, transverse stripe; lower paraocular area withirregular triangular yellow mark along the eye and up to level of antennal socket; supraclypealarea unmarked. Pronotum with transverse collar stripe yellow, interrupted medially; lobesyellow. Legs dark brown except fore tibia with pale area; tibiae with small, basal yellowspots (lacking in Kauai specimens). Wings slightly smoky.

Perkins’ collections. Hawaii: Kilauea (BMNH).Recent collections. Hawaii: Kona Forest Unit of Hakalau National Wildlife Refuge,

South Boundary Rd., 5200 ft; camp, 5300 ft. Maui?: W. Maui, Honokohau, 900 m, 26-II-1972, J. L. Gressitt (BPBM). Oahu: Manana Trail, 1800 ft. Kauai: Awaawapuhi Trail,3800 ft; Na Pali Kona Forest Reserve, 4100 ft.

Flower records. Cheirodendron, Claoxylon, Metrosideros, Myoporum, Scaevola procera.Remarks. Both Perkins (1911:723) and Fullaway (1918:395) commented on the possible

synonymy of H. specularis, described from the female taken at Kilauea, and H.homoeochromus, described later from the male taken at Kilauea. The only type available isthat of H. specularis. Hylaeus homoeochromus is treated here as a junior synonym becausemales and females that match the descriptions of the respective taxa have been collectedtogether by D. T. Fullaway and KM, and have been genetically associated. Two femalesfrom Maui appear to be this species, although the impunctate area next to the lateral ocellusis not as shiny as in other specimens. Regardless of their identity, the recent discovery ofthis species on Kauai and Oahu means that it almost certainly inhabits Maui Nui as well.Males from Kauai have consistently broader, more curved, and more pointed gonoforcepsthan Hawaii specimens. Males from Oahu were not available to compare. Although foundon many islands, it appears to be local and rare on all.

See also remarks for H. crabronoides and H. dumetorum.References. Perkins, 1907b: 97 (record: Hawaii, Kilauea). Hylaeus homoeochromus:

Meade-Waldo, 1923:35 (list).

Hylaeus (Nesoprosopis) sphecodoides (Perkins)(Fig. 64)

Nesoprosopis sphecodoides Perkins, 1899:105-106 (♂♀; cleptoparasite of H. difficilis).Hylaeus (Nesoprosopis) sphecodoides, Meade-Waldo, 1923:34 (list).Nesoprosopis sphecodoides, Daly, 1994:119 (lectotype ♂ BMNH 17a-137b; Hawaii,

Kau, 4000 ft; wing length, 4.5 mm).


DIAGNOSIS. Medium-sized cleptoparasitic bees with clear to slightly smoky wings andwith or without red basal terga. Male face with three separate yellow marks (clypeus,paraocular areas), malar space very short, scape slightly dilated. Female face black or darkreddish, paraocular areas not marked, clypeus emarginate, frons with indistinct punctures,fore tarsus with straight hair. See discussion of cleptoparasites under H. hilaris.

Distribution. Hawaii; coastal to montane dry shrubland and dry to mesic forest.Male. Measurements: Malar space, 3; eye, 120:106:102:72; clypeus, 56×61; frons,


Head. Malar space very short (ratio about 0.2-0.3 DMO). Upper paraocular area withindistinct, round pits 1-2 pit widths apart, may be obscured by minute, parallel wrinkles;frons with pits more distinct medially, smaller, and contiguous; middle dull with dense pits;with or without fine frontal line; large impunctate areas above antennal sockets. Lowerparaocular depression absent. Frontoclypeal sulcus straight. Scape in frontal view slightlydilated, width half length, ratio about 0.5; lateral edge in frontal view nearly straight; slightlyarched in lateral view; no groove on underside; with medium-length, white, weakly plumosehair. Vertex hair 6-10, white, plumose.

Mesosoma. Scutum with pits about 1 pit width apart or less, ground slightly lustrous;hair short (6-8), white, plumose. Scutellum similar. Lateral carina of front coxa lamellate,produced into a short spine (Fig. 8D). Mesepisternum with indistinct, shallow pits about 1pit width apart, and long, white, plumose hair; ground nearly smooth, slightly lustrous.Hypoepimeron impunctate or with weak punctation, smooth, slightly lustrous, slightlyswollen. Metanotum slightly lustrous. Basal area of propodeum lineately or reticulatelyrugose to poorly defined brow, shiny between rugae at front; brow minutely wrinkled, dull.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, very short,pale, appressed hairs spaced more than a hair length apart. T7 hair short, yellowish whiteto pale brown, prostrate.

Terminalia. Lateral process of S7 parallel-sided, length greater than width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with short, triangular tips. Apicesof gonoforceps subequal to penis valves.

Color. Labrum and mandible blackish brown to black. Face with three separate marks:clypeus with large, central yellow spot surrounded by dark cuticle; lower paraocular areamostly yellow below antennal socket or extended in a narrowing stripe to just above socket;supraclypeal area and scape unmarked. Pronotum unmarked. Head and mesosoma black;metasoma with basal 1-2 segments reddish brown and apical segments dark brown orentire metasoma dark blackish brown to light brown. Legs dark brown to blackish brown,unmarked except pale area on fore tibia and pale brown fore basitarsus; tibiae and tarsimay be reddish brown. Wings clear to slightly smoky.

Female. Head. Malar space very short (ratio about 0.2 DMO). Upper paraocular areawith shallow, indistinct pits about 1 pit width apart, ground with or without minute parallelwrinkles; in some bees, frons with pits smaller medially, closer, and obscure at middle, inother bees pits distinct near middle; with or without a fine frontal line; large impunctate


areas above antennal sockets. Apex of clypeus deeply emarginate. Dorsal end of facialfoveae not or barely extended to level of lateral ocelli. Vertex hair 6-10, white, plumose.

Mesosoma. Fore tarsus with hair short, straight (see Fig. 8B). Scutum with pits about 1pit width apart, ground slightly granular, making pits less distinct, dull to slightly lustrous;hair short (2-6), white, plumose. Lateral carina of front coxa lamellate, produced into ashort spine. Mesepisternum with indistinct or distinct, shallow pits about 1 pit width apart,and yellowish white, plumose hair; ground nearly smooth to minutely wrinkled, slightlylustrous. Hypoepimeron impunctate, ground with minute, parallel, longitudinal wrinkles;nearly flat. Metanotum dull. Basal area of propodeum lineately rugose to poorly definedbrow, shiny between rugae at front; brow minutely wrinkled, dull.

0.5 mm

A B C D

FE

1.0 mm

Fig. 64. Hylaeus sphecodoides, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous; with fine, extremelyshort, pale, appressed hairs many hair lengths apart. T6 hair medium to long, light brown,prostrate.

Color. Head and mesosoma dark reddish brown to black; metasoma with basal 2 segmentssomewhat pale or reddish brown and apical segments dark to light brown. Face and pronotumunmarked. Legs brown to dark brown except pale area on fore tibia; unmarked. Wingsclear to slightly smoky.

Perkins’ collections. Hawaii: Kau, 4000 ft (BMNH, BPBM, OUMNH); Kilauea (BPBM,OUMNH); Kona, 4000 ft (BMNH, BPBM).

Recent collections. Hawaii: Kipuka Nene, 2900 ft; Keauhou Ranch, 4500 ft; KipukaAlala, 6100 ft; South Point, coast.

Flower records. None.Remarks. Hylaeus sphecodoides and H. volatilis have melanic forms with no red on the

metasomal terga and, in H. volatilis, entirely black with no yellow marks (see remarks forthat species). See also discussion of cleptoparasites under H. hilaris.

References. Perkins, 1907b: 97 (record: Hawaii, Kilauea); 1910:603 (key). Suehiro,1986:61 (record: Hawaii, Kona).

Hylaeus (Nesoprosopis) takumiae Magnacca and Daly, new species(Fig. 65)

DIAGNOSIS. Medium-sized bees with an unusual shiny area between the lateral ocellusand eye, unusually long malar space, and clear wings. Male face without marks or only asubapical yellow spot on clypeus, and unusual extremely dilated scape. Female black andunmarked. See remarks.

Distribution. Maui; subalpine dry shrubland.Male. Measurements: Malar space, 12; eye, 120:118:105:76; clypeus, 64×69; frons,


Head. Malar space long (ratio about 0.8 DMO). Upper paraocular area with sparse,scattered, nearly round pits and minutely striate, lustrous ground; frons to middle distinctlydifferent with small dense pits, distorted, and contiguous, with dull ground; denselypunctured area extended ventrally to antennal sockets; frontal line mostly obscured bydense pits; conspicuous impunctate, shiny area between lateral ocelli and eye. Lowerparaocular area with distinct depression, impunctate, and with oblique, minute, parallelwrinkles. Clypeus with shallow pits 1 pit width apart. Upper clypeus distinctly duller thanlower. Frontoclypeal sulcus arched; supraclypeal area as wide as long; clypeus protuberant,subapically convex in lateral view. Frons with long, dark brown hair. Scape in frontal viewextremely dilated, cordiform, width greater than length, ratio about 1.1; lateral edge infrontal view strongly convex; moderately arched in lateral view, excavated underneath andwith no groove, with maximum arch near middle of scape; with long, dark brown, simplehairs. Vertex hair 24-40, dark brown, mostly simple.


Mesosoma. Scutum with distinct, round pits about 1-3 pit widths apart, ground smoothlycoriaceous, lustrous; hair very long (20-30 or more), brown, weakly plumose. Scutellumwith pits less distinct, ground lustrous. Mesepisternum with indistinct pits about 1-2 pitwidths apart, and long, brownish white to brown, plumose hair; ground minutely roughened,lustrous. Hypoepimeron obscurely punctured in lower half, swollen, lustrous. Metanotumminutely roughened, slightly lustrous. Basal area of propodeum reticulately rugose orareolate to brow, shiny between rugae; densely roughened at and below brow, but lustrous.

Metasoma. Disk of T2 without distinct punctation, smooth, shiny, with short, pale hairsmore than a hair length apart. T7 hair long, relatively sparse, dark brown, erect.

Terminalia. Lateral process of S7 tapered, length subequal to width of hemisternite.Median process of S8 somewhat swollen in basal half, angled at midlength, slender in distalhalf, apex bifurcate with moderately long, narrow tips. Apices of gonoforceps subequal topenis valves.

Color. Head and mesosoma black; metasoma dark reddish brown to black. Labrum andmandible black. Face usually unmarked (one bee with small, subapical yellow mark onclypeus); scape unmarked; antennal flagellum reddish yellow to pale yellow beneath,prothorax unmarked. Legs dark brown to black, unmarked except pale area on fore tibia.Wings clear, iridescent.

Female. Head. Malar space long (ratio about 0.8). Apex of clypeus shallowly emarginate.Dorsal end of facial fovea extended to level of lateral ocellus. Upper paraocular area withnearly round pits less than 1 pit width apart, on frons becoming medially smaller, distorted,contiguous; frontal line obscured by dense pits; conspicuous impunctate, shiny area betweenlateral ocelli and eye. Vertex hair 20-40, dark brown, weakly plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with distinct pits about 1-3pit widths apart, ground smoothly coriaceous, lustrous; hair very long (14-30), brown,weakly plumose. Scutellum with pits less distinct, ground lustrous. Mesepisternum withindistinct pits about 1-2 pit widths apart, and yellowish white, plumose hair; ground minutelyroughened, lustrous. Hypoepimeron obscurely punctured in lower half, swollen, lustrous.Metanotum minutely roughened, slightly lustrous. Basal area of propodeum reticulatelyrugose or areolate to brow, shiny between rugae.

Metasoma. Disk of T2 without distinct punctation, smoothly coriaceous, lustrous, withfine, short, pale hairs spaced more than a hair length apart. T6 hair long, dark brown, erect.

Color. Face, scape, and pronotum unmarked. Legs dark brown to black, with or withoutobscure pale area on fore tibia. Wings clear.

Holotype ♂ labeled “Maui, Haleakala National Park, Haleakala West slope, 2865 m,13-Jun-1997, R. Takumi” (CUIC).

Allotype ♀ labeled “Maui Co., Haleakala NP, malaise 2500 m, 20-Jun to 2-Jul-1991, R.Takumi” (CUIC).

Paratypes. Maui. Haleakala NP, NW slope between Leleiwi Trail to overlook andHalemauu Trail, 2500 m, malaise trap, 1♂ 4♀ 14-20-VI, 1♂ 12♀ 20-VI to 2-VII, 1♂ 5♀2-5-VII, 5♀ 5-12-VII, 7♀ 12-24-VII-1991 (RT) (CUIC). Sliding Sands Tr., 2600-3000m, 4♂ 17-VI-1986, F. G. Howarth; Puu Maile, 7500 ft, 2♂ 21-VI-1976, J. W. Beardsley;Kapalaoa, 7280 ft, 2♂ 18-VI-1976, J. W. Beardsley; S. of Holua, 7300 ft, 1♂ 4-VI-1977,A. Hara; West Kaupo Gap, 5700 ft, 1♂ 7-VI-1977, G. Teves (BPBM).


Flower records. Coprosma montana, Styphelia.Etymology. The species epithet takumiae is in honor of the collector, Raina Takumi.Remarks. This species is an inhabitant of the subalpine shrubland of East Maui. It has

been collected on the outer northwest slopes of Haleakala as well as within the summitcrater at Kapalaoa Cabin. It is an unusually fast flier, much more so than the more commonH. nivicola, and this may be part of the reason it was not collected earlier.

Common species of similar body size that occur in this general area are H. difficilis, H.nivicola, H. volcanicus, and the cleptoparasite H. volatilis. Both sexes of H. takumiae aredistinguished from these species by an impunctate, polished, shiny area between the lateralocellus and eye. Both sexes of the other listed species have the ocellocular areas minutelytextured and usually dull. The malar space of H. takumiae is long, with a ratio of about 0.8DMO and is similar to that of H. volcanicus (the malar space of H. nivicola is longer and

Fig. 65. Hylaeus takumiae, new species, male. A. Seventh sternum, ventral view. B. Eighth sternum,ventral view, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E.Face with left scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E.

0.5 mm

A B C D

F

1.0 mm

E


the malar space of the other species is usually shorter). The females of H. takumiae havethe basal area of the propodeum reticulately rugose, whereas those of H. volcanicus havethe basal area lineately rugose. Males of H. takumiae are distinguished from the otherlisted species by the extremely dilated, cordiform scape (all other species listed have muchless dilated scapes). However, H. satelles, not recently collected in this area, has a similarlydilated scape in the male and shiny areas next to the lateral ocelli in both sexes. Both sexesof H. satelles are larger in body size, with punctured metasomal terga, dark wings, andshort malar areas.

Hylaeus (Nesoprosopis) unicus (Perkins)(Fig. 66)

Nesoprosopis unica Perkins, 1899:88 (♂; holotype ♂ BMNH 17a 123; Oahu, HonoluluMt., 2000 ft).

Hylaeus (Nesoprosopis) unicus, Meade-Waldo, 1923:34 (list).Nesoprosopis unica, Daly, 1994:116 (holotype confirmed).Nesoprosopis laticeps Perkins, 1899:88-89 (♂♀; Pl. 2, Fig. 15) (new synonymy).Hylaeus (Nesoprosopis) binominatus Meade-Waldo, 1923:34 (list; replacement name

for Nesoprosopis laticeps Perkins, a junior homonym of Hylaeus laticeps Morawitz)(new synonymy).

Prosopis avara Warncke, 1970:747 (unwarranted replacement name for Hylaeus laticepsPerkins) (new synonymy)

Nesoprosopis laticeps, Daly, 1994:117 (lectotype ♂ BMNH 17a 124b; Molokai, Mts.,4000 ft; wing length, 5.4 mm).

DIAGNOSIS. Medium-sized bees with clear to smoky wings and unique, extremely widesupraclypeal area. Male face black or with yellow mark in lower paraocular area or alsoclypeus; scape moderately dilated and with lateral margin concave and unusual maximumarch near apex. Female black and unmarked or with small yellow marks on paraocularareas and sides of pronotal collar.

Distribution. Maui (new record), Lanai, Molokai, and Oahu; montane wet and mesicforest.

Male. Measurements: Malar space, 3; eye, 108:114:108:66; clypeus, 55×65; frons,15×34:74; interalveolar, 25:22:46:52, D 22; interocellar, 31:34:22:31, D 18; scape, 39×28;wing length, 4.7 mm (H. unicus holotype).

Head. Malar space very short to short (ratio about 0.2-0.3 DMO). Upper paraoculararea and frons with nearly round pits about 1 pit width apart, pits closer but still nearlyround near middle, ground minutely wrinkled and slightly lustrous between pits; middlearea between median ocellus and acme with faint frontal line or only a few fine longitudinalwrinkles. No distinct impunctate area above antennal socket. Lower paraocular depressiondistinct, but shallow. Supraclypeal area much wider than long, W/L ratios 1.8-2.6.Frontoclypeal sulcus arched. Scape in frontal view slightly to moderately dilated, widthmore than half length, ratios about 0.6-0.8, widest near apex; lateral edge in frontal viewstraight or slightly concave and slightly angled at apex; strongly arched in lateral view with


0.5 mm

A B C D

E F

1.0 mm

Fig. 66. Hylaeus unicus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.

maximum arch nearer apex of scape, apex expanded; no groove on underside; with long,brown, mostly simple hair. Vertex hair 16-30, brown, mostly simple.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground dull; hair medium to long (8-16), brown, mostly simple. Scutellum with very slightly larger pits, ground slightly lustrousin contrast to scutum. Mesepisternum with indistinct, shallow, round pits, about 1 pit widthapart, and long, yellowish white, weakly plumose hair; ground minutely coriaceous, slightlylustrous. Hypoepimeron impunctate, lustrous, slightly swollen. Metanotum dull. Basal areaof propodeum with sparse, lineate to nearly reticulate rugae extended about halfway tobrow; brow smoothly coriaceous and lustrous to shiny.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart or less. T7 hair long, brown, erect.


Terminalia. Lateral process of S7 tapered, with length subequal to width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with broad tips. Apices ofgonoforceps subequal to penis valves.

Color. Head and mesosoma black; metasoma dark brown to blackish brown. Labrumand mandible blackish brown to black. Facial marks highly variable: unmarked (Maui Nui);or with small triangular mark in lower paraocular area below level of antennal socket, withor without a subapical clypeal mark (Maui Nui); or with lower paraocular area largely orentirely yellow below antennal socket and clypeal mark large, up to entire apical half ofclypeus except margins (Oahu). Supraclypeal area and scape unmarked. Pronotum unmarkedor with small yellow marks at sides of collar. Legs black, fore tibia and apex of fore femurwith pale area; small, obscure yellow spot at base of fore tibia, up to half length in somespecimens; small subapical yellow mark on mid tibia present in some specimens; obscureyellowish marks at bases of mid and hind tibiae present in some specimens; basitarsi withor without yellow marks. S6 pale yellow. Wings clear to smoky.

Female. Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area andfrons with nearly round pits about 1 pit width apart, pits mostly evenly spaced and nearlyround even near the middle, ground minutely wrinkled and slightly lustrous between pits;no impunctate area above antennal socket; faint frontal line. Apex of clypeus shallowlyemarginate. Dorsal end of facial fovea extended to level of lateral ocellus. Vertex hair 14-20(-24), brown, weakly plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with distinct pits, 1-2 pitwidths apart, ground dull; hair medium-length (6-12), brown, simple. Scutellum with pitsslightly larger, ground lustrous in contrast to scutum. Mesepisternum with indistinct, shallow,round pits, about 1-2 pit widths apart, and long, yellowish white, weakly plumose hair;ground minutely coriaceous, slightly lustrous. Hypoepimeron impunctate, lustrous, slightlyswollen. Metanotum dull. Basal area of propodeum weakly lineately rugose less than halfwayto brow; brow smoothly coriaceous and lustrous.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, lustrous, with fine, pale,appressed hairs spaced about one hair length apart or less. T6 hair long, dark brown toblack, erect.

Color. Head and mesosoma black; metasoma and legs blackish brown; unmarked orwith small yellow stripes on lower paraocular areas and sides of pronotal collar. Wingssmoky.

Perkins’ collections. Nesoprosopis laticeps: Lanai: Halepaakai, 3000 ft (BPBM); KoeleMts., <3000 ft (BPBM), 3000 ft (BMNH, BPBM, OUMNH); “Lanai,” 2000 ft (BMNH).Molokai: “Molokai,” 4500 ft (OUMNH); Molokai Mts., 4000 ft (BMNH). N. unica: Oahu:Honolulu, 2000 ft (BMNH); Pauoa Valley, head of (BPBM); Waialua, 1800 ft (BPBM).

Recent collections. Maui: Puu Kukui Rd., 2300 ft; Haelaau Cabin, 3000 ft; Puu KukuiTr., 3700 ft. Lanai: Munro Trail, 3000 ft. Molokai: Kawela intake, 3600 ft; Pepeopaeboardwalk, 4000 ft; Kamakou Rd., 3200 ft. Oahu: Puu Kaua, 2600 ft; Wiliwilinui Trail,2000 ft; Papali Ridge, 1500 ft; Koolau Summit Trail, 2300 ft; Mt. Kaala, 4000 ft.

Flower records. Acacia, Cheirodendron, Dubautia laxa, Metrosideros, Psychotriafauriei, Scaevola chamissoniana, S. gaudichaudiana, Tetraplasandra oahuensis.


Remarks. Fullaway (1918:394) pointed out the possible synonymy of H. kauaiensiswith H. unicus and H. binominatus, known then as N. laticeps. Although clearly allied, H.kauaiensis is structurally distinct from bees referred to the other two names. However, H.unicus described from Oahu and H. binominatus from Lanai and Molokai are consideredhere to apply to one species under the name H. unicus. This name has page precedence andis not involved in homonymy. Males are closely similar in the extreme width of thesupraclypeal area; shape, dilation, and arch of the scape; propodeal sculpture; and terminalia.On Oahu males are less melanic: a large or small spot on the clypeus, large triangularmarks on the paraocular areas, marks at the sides of the pronotal collar, yellowish marks atthe bases of the mid and hind tibiae and tarsi, and wings slightly smoky (the figure is of anOahu specimen). Some females from Oahu have small yellow stripes on the paraocularareas, marks at the sides of the pronotal collar, and smoky wings. On Maui and Lanai theface of males is entirely dark or with triangular marks or traces of yellow in the paraocularareas; no marks on the pronotum or legs; and smoky wings. Females are unmarked andwith smoky wings.

References. Perkins, 1910:603 (key). Swezey, 1913:198 (record: Oahu, Punaluu);1922:14 (nest: in Pipturus twig; parasitoid: Eupelmus near E. euprepes; record: Oahu,Tantalus); 1954:167 (nest: in Pipturus; record: Oahu, Tantalus). Nesoprosopis laticeps:Perkins, 1910:603 (key); 1913:79 (biogeography).

Hylaeus (Nesoprosopis) volatilis (F. Smith)(Fig. 67)

Prosopis volatilis F. Smith, 1879a: 683-684; 1879b: 21-22 (♂; holotype ♂ BMNH 17a152, “Kauai,” but see Blackburn and Cameron [1886:148], who stated that Oahu iscorrect type locality and Perkins [1899:105] does not record collections from Kauai;wing length, 4.2 mm).

Nesoprosopis volatilis, Perkins, 1899:105 (♂♀; cleptoparasite of N. assimulansoahuensis, N. facilis, N. difficilis).

Hylaeus (Nesoprosopis) volatilis, Meade-Waldo, 1923:34 (list).Hylaeus volatilis Houston, 1981:50 (resolution of error involving the type label of N.

volatilis on the wrong specimen).Nesoprosopis volatilis, Daly, 1994:120 (holotype confirmed).

DIAGNOSIS. Medium-sized cleptoparasitic bees with clear to slightly smoky wings andwith or without red basal terga. Male face marks variable from no marks to entire lowerface yellow (clypeus, supraclypeal area, and paraocular areas) and extended above antennalsockets in narrowing stripe, scape slightly dilated. Female face black or dark reddish,paraocular areas not marked, clypeus emarginate, frons with distinct punctures, fore tarsuswith straight hairs. See discussion of cleptoparasites under H. hilaris.

Distribution. Maui, Lanai, Molokai, and Oahu; coastal to subalpine dry shrubland. Recentcollections only from Maui.


Male. Measurements: Malar space, 4; eye, 146:125:120:79; clypeus, 70×67; frons,24×26:100; interalveolar, 18:29:61:69, D 18; interocellar, 28:36:28:38, D 15; scape, 51×29;wing length, 4.9 mm (Oahu, Waianae, 2000 ft, BPBM 3639).

Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area with round pits1-2 pit widths apart, ground with minute parallel wrinkles (bees with stronger wrinkleshave indistinct pits); frons with pits becoming contiguous medially and smaller, groundsmoother, middle not distinguished by wrinkles or a frontal line; frons slightly lustrousoverall; large, impunctate, lustrous area above antennal sockets. General area of vertexand frons slightly swollen and gently convex. Lower paraocular depression absent.Frontoclypeal sulcus nearly straight to slightly arched. Scape in frontal view slightly dilated,width a little more than half length, ratio about 0.6; lateral edge in frontal view straight toslightly concave; slightly arched in lateral view; no groove on underside; with short, yellowishwhite, mixed simple and plumose hair. Vertex hair 8-14, white, plumose.

Mesosoma. Scutum with pits 1-2 pit widths apart, ground smoothly coriaceous, slightlylustrous; hair short (6-8), white, plumose. Scutellum with slightly larger pits, slightly lustrous.Lateral carina of front coxa lamellate, produced into a short spine. Mesepisternum withdistinct, round pits about 1 pit width apart, and long, white, plumose hair; ground smoothlycoriaceous, slightly lustrous. Hypoepimeron impunctate, smooth, swollen. Metanotum dull.Basal area of propodeum short, rather flat and without definite brow, roughened overallwith lineate rugae at front, becoming more indistinct and reticulate distad, shiny betweenrugae at front.

Metasoma. Disk of T2 impunctate, smoothly coriaceous, shiny; with fine, short, pale,appressed hairs spaced about one hair length apart; without bands of hairs. T7 hair short,brown, appressed.

Terminalia. Lateral process of S7 parallel-sided, length greater than width of hemisternite.Median process of S8 moderately dilated, apex bifurcate with short, triangular tips. Apicesof gonoforceps subequal to penis valves.

Color. Labrum and mandible yellowish brown to brown, or black in melanic bees. Facevariable, with a single large spot, three separated marks, two marks only on paraocularareas, or unmarked: clypeus entirely yellow to unmarked; paraocular area entirely yellowextended in narrow vitta to well above antennal socket, or only small spot, or unmarked;supraclypeal area variable, from unmarked to yellow to acme; scape with yellowish lateralmargin or unmarked. Metasoma with two basal segments and base of third segment redand apical segments dark brown, or metasoma entirely black in melanic bees. Pronotumunmarked. Legs dark brown to black except: fore tibia with pale area with or withoutyellowish, dorsal stripe; fore basitarsus yellowish brown to light brown, or black in melanicbees; all tibiae with reddish areas at base. Wings clear to slightly smoky.

Female. Head. Malar space short (ratio about 0.3 DMO). Upper paraocular area andfrons with round pits 1-2 pit widths apart; bees from Oahu and some from Maui haveground with minute, parallel, dull wrinkles and indistinct pits at sides and near the middle,pits more distinct in the intervening area; some bees from Maui have distinct pits withlustrous, smooth ground across areas; with or without a fine frontal line; large, impunctate,


lustrous area above antennal sockets. Apex of clypeus deeply emarginate. Dorsal end offacial foveae not extended to level of lateral ocelli. Vertex hair 4-8, white, plumose.

Mesosoma. Fore tarsus with hairs short, straight (see Fig. 8B). Scutum with pits 1-2 pitwidths apart, ground smoothly coriaceous, slightly lustrous; hair very short (2-4), white,simple. Lateral carina of front coxa lamellate, produced into a distinct spine. Mesepisternumwith round pits about 1 pit width apart, and medium-length, yellowish white, plumose hair;pits distinct in some bees with smooth ground, in other bees pits partly obscured by minutelyroughened, wrinkled or striate ground; dull to slightly lustrous. Hypoepimeron impunctate,striate, slightly swollen. Metanotum dull. Basal area of propodeum short, rather flat andwithout definite brow, roughened overall with lineate rugae at front, becoming less distinctand reticulate distad, shiny between rugae at front.

0.5 mm

A B C D

E F

1.0 mm

Fig. 67. Hylaeus volatilis, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventral view,and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face with leftscape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescence shownby dashed lines in B-E.


Metasoma. Disk of T2 impunctate, smoothly coriaceous, shiny; with fine, extremelyshort, pale, appressed hairs many hair lengths apart; without bands of hairs. T6 hair long,light brown or brown, appressed.

Color. Head and mesosoma black; metasoma with basal two or three segments red andapical segments reddish brown or entirely dark reddish brown (nearly black) in melanicbees. Face marks: clypeus, lower paraocular areas, supraclypeal area, mandibles, and lateralmargin of scape an obscure reddish color, or face entirely black in melanic bees. Pronotumunmarked. Legs without yellow marks, dark brown to black except: fore tibia with palearea, fore legs yellowish brown, brown, or black, all tarsi reddish brown. Wings clear toslightly smoky.

Perkins’ collections. Maui: E side Haleakala, 7-10,000 ft (OUMNH); Iao Valley(BPBM), far up (BPBM); “Maui” (BPBM); Wailuku (BMNH, BPBM), sandy isthmus(OUMNH). Lanai: Lanai, 2000 ft (BMNH). Molokai: Kaulawai (BPBM); Molokai, farwest end (BPBM); Molokai Mts., 3000 ft (BMNH, BPBM, OUMNH); Molokai, plains(BPBM). Oahu: Honolulu (BPBM), 2000 ft (BMNH); Honolulu Mts., 1500 ft (BPBM);Waianae (OUMNH), 2000 ft (BPBM); Waianae, Makana coast (BPBM); Waianae Mts,2000 ft (BMNH, BPBM), 2-3000 ft (BPBM).

Recent collections. Maui: Haleakala N. P. Visitor Center, 2975 m; above Hosmer Grove,2100 m; Kalahaku, 2835 m; Kuiki, 2170 m; Kaluaiki, 2225 m; Laie, 2040 m; Lauulu Trail.,1955 m; Oili Puu, 2040 m; Paliku, 1830-1950 m; Puu Maile, 2195 m; Waikane, 2010 m;Halemauu Tr., 7800-8000 ft; Waikamoi Preserve boardwalk, 6400 ft; Puu Keokea, 7100ft; Kaupo Tr., 5500 ft; Lahainaluna, 1800 ft.

Flower records. Styphelia.Remarks. Hylaeus volatilis exhibits considerable variation in body size and the pattern

of yellow marks. Perkins (1913:lxxix) noted that H. volatilis ranges from coasts to 9000 ftin the mountains. He also observed that cleptoparasites may vary in size according to thesize of the host’s provisions and could be exceptionally large when the hosts were large(1913:lxxxi; see also Michener 2000:30). Collections of this species on Haleakala yieldedbees ranging in size (wing length) from 4.0 to 5.1 mm and with one male at 3.5 mm. AtHaleakala, the common probable hosts are H. difficilis, H. nivicola, H. takumiae, and H.volcanicus, all of which are about the same size. The cleptoparasites in this area exhibitvariation in the extent of melanism. Males range from individuals with yellow marks andred basal terga to melanic individuals that are almost entirely dark brown to black. Among41 males collected during 1991-1997, the faces of 6 (15%) are marked with at least someyellow on the paraocular areas (extended up along the eye in well-defined, narrow stripeabove the level of the antennal socket or more frequently reduced to an obscure spot ateach side); less commonly also on the clypeus as a central spot; and with the basal 1 to 3metasomal segments partly red. Intermediate bees (16, or 39%) have the face black ornearly so and with at least the basal metasomal tergum partly red. Melanic bees (19, or46%) have the face and metasoma black; many retain a small red spot laterally on T1.Regardless of the degree of melanism, males have the following features in common:subglabrous metasomal terga without pubescent margins; basal half of the propodeal areawith coarse rugae, mainly linear and longitudinal, with the ground flat and shiny between


rugae; apical half of the propodeal area less shiny with finely reticulate rugae; S7 with longlateral processes; median process of the S8 moderately dilated and pubescent at the flexureand the slender apex bifid with short, pubescent tips; and similar male genitalia with long,apical, weakly plumose hair. Among 36 females collected during 1991-1997, 7 (19%) havethe clypeus, and rarely also the lower paraocular areas to the level of the antennae, adiffuse red color; the mandibles, scape, and fore tibia (both in frontal view) and up to 3basal metasomal segments are partly red. Intermediate females (28, or 78%) have the faceand scape black or nearly so, reddish fore tibiae, and up to 2 basal metasomal segmentsred. The single melanic female has only the fore tibiae reddish. All females have an emarginateclypeus, and as in the males, the coarse sculpture on the basal half of the propodeum andsubglabrous metasoma.

See discussion of cleptoparasites under H. hilaris.References. Blackburn and Kirby, 1880:85 (record). Blackburn and Cameron, 1886:148

(record, key); 1887:207-208 (record, key). Dalla Torre, 1896:35 (list). Perkins, 1907a: 43(record: Oahu, Tantalus); 1910:604 (key); 1913:79 (biogeography). Michener, 1965:129(list; see Houston, 1981:50). Beardsley, 1980:40 (record: Maui, Haleakala).

Hylaeus (Nesoprosopis) volcanicus (Perkins)(Fig. 68)

Nesoprosopis volcanica Perkins, 1899:83 (♂♀; Pl. 2, Fig. 12).Hylaeus (Nesoprosopis) volcanicus, Meade-Waldo, 1923:33 (list).Nesoprosopis volcanica, Daly, 1994:119 (lectotype ♂ BMNH 17a 135a; Hawaii, Kau,


DIAGNOSIS. Medium-sized bees with clear wings. Male with long malar space, scapeslightly dilated, face with oval yellow mark (clypeus, supraclypeal area, lower paraocularareas) divided by fine black sulci, lateral process of S7 narrow, and median process of S8not dilated and with long narrow tips. Female with medium to long malar space; black andunmarked. For similar species, see remarks for H. nivicola and H. difficilis.

Distribution. Hawaii and Maui; dry montane and subalpine shrubland, usually on recentlava flows.

Male. Measurements: Malar space, 12; eye, 131:118:108:77; clypeus, 69×69; frons,26×24:91; interalveolar, 15:23:54:61, D 22; interocellar, 31:33:23:33, D 15; scape, 46×22;wing length, 4.6 mm (Hawaii, Kau, 4000 ft, BPBM 3653).

Head. Malar space long (ratio about 0.8-1.0 DMO). Upper paraocular area and fronswith nearly round pits less than 1 pit width apart, ground minutely wrinkled, dull; minute,parallel wrinkles down along the eye; median, impunctate, slightly lustrous area aboveantennal sockets, with a minute, central, elliptical hole in some bees. Lower paraoculardepression distinct. Frontoclypeal sulcus arched. Scape in frontal view slightly dilated,width half length, ratio about 0.5; lateral edge in frontal view nearly straight or slightlyconcave; slightly arched in lateral view with maximum arch near middle of scape; no grooveon underside; with medium-length, yellowish white, mixed simple to weakly plumose hair.Vertex hair 16-20, brown, simple to weakly plumose.


Mesosoma. Scutum with indistinct pits 1-2 pit widths apart, ground minutely wrinkled,dull to slightly lustrous; hair long (16-30), yellowish white, mixed simple to plumose.Mesepisternum with pits indistinct, round, shallow, 1 pit width apart or less; with long,white, plumose hair; ground minutely roughened, wrinkled, or striate; dull to slightly lustrous.Hypoepimeron impunctate, smooth to weakly striate. Metanotum dull. Basal area ofpropodeum lineately rugose to brow or beyond, or lineate at front to reticulate at brow orbeyond, shiny between rugae at front; brow granularly coriaceous or with minute, reticulaterugae, dull to slightly lustrous.

Metasoma. Disk of T2 impunctate, lustrous; fine, pale, appressed hairs spaced morethan a hair length apart. T7 hair short, golden, appressed.

Terminalia. Lateral process of S7 tapered, with pale apex, length subequal to or greaterthan width of hemisternite. Median process of S8 linear, apex bifurcate with long, filamentoustips. Apices of gonoforceps subequal to penis valves.

Color. Body black. Labrum and mandible blackish brown to black. Face with a singlelarge spot: clypeus entirely yellow or nearly so; lower paraocular area variable, stripe ofyellow below antennal socket and along sulcus or area unmarked; supraclypeal area toacme yellow or unmarked; scape unmarked. Pronotum unmarked. Legs dark brown toblack except pale area on fore tibia and, at most, obscure basal yellowish spots on tibiae.Wings clear.

Female. Head. Malar space medium to long (ratio about 0.6-0.9 DMO). Upper paraoculararea and frons with round, shallow pits 1 pit width apart or less; ground minutely wrinkled,slightly lustrous; minute longitudinal wrinkles at middle (sometimes obscuring pits) or finefrontal line sometimes present. Apex of clypeus shallowly emarginate. Dorsal end of facialfovea extended to level of lateral ocellus. Vertex hair 16-20, brown, plumose.

Mesosoma. Fore tarsus with hairs apically curved. Scutum with distinct pits 1-2 pitwidths apart, ground minutely wrinkled, slightly lustrous; hair long (12-20), pale brown,plumose. Scutellum with pits slightly farther apart, ground smooth, lustrous. Mesepisternumwith pits relatively distinct compared to male (may be indistinct if ground highly roughened),round, shallow, 1 pit width apart or less; with medium-length, yellowish white, plumosehair; ground minutely wrinkled or striate, dull to slightly lustrous. Hypoepimeron impunctate,smooth to weakly striate; slightly swollen. Metanotum dull. Basal area of propodeumlineately rugose to brow or beyond, or lineate at front to reticulate at brow or beyond,shiny between rugae at front; brow granularly coriaceous or with minute reticulate rugae,dull to slightly lustrous.

Metasoma. Disk of T2 impunctate, lustrous, with fine, short, pale, appressed hairs spacedmore than a hair length apart. T6 hair light brown to golden, medium-length, appressed.

Color. Body and legs black; unmarked. Wings clear.Perkins’ collections. Hawaii: Kau, 4000 ft (BMNH, BPBM); Kilauea (BPBM); 4000

ft (OUMNH); Kona, 5000 ft (BMNH). Maui: Haleakala, 9000 ft (OUMNH); crater(BMNH, BPBM).

Recent collections. Hawaii: Footprints, 900 m; Kipuka Nene, 2900 ft; Napau Trail,3000 ft; Devastation Trail, 3700 ft; Kipuka Puaulu, 4000 ft; Kulani Cone, 5100 ft; KipukaMaunaiu, 1890 m; Mauna Loa tr. 16, 6000 ft; Hualalai summit, 8000 ft. Maui: ScienceCity, 3010 m; near Haleakala N. P. Visitor Center, 9750 ft; Laie flats, 2040 m; Oili Puu,


2020 m; Paliku, 1950 m; Sliding Sands Tr., 2500 m; Waikane, 2010 m; Holua Cabin, 7200ft; Halemauu Tr., 7800-8000 ft; Lauulu Tr., 8000 ft; Kalapawili Ridge, 7400 ft; Kaupo Tr.,5500 ft; Kilohana, 2800 m; Kipahulu Valley, Mauka Ridge, 1980 m; Puu Keokea, 7100 ft;Lihau, 4000 ft.

Flower records. Coprosma sp., Dodonaea, Dubautia menziesii, Dubautia ×Argyroxiphium, Metrosideros, Myoporum, Sophora, Styphelia.

Remarks. See remarks for H. nivicola and H. difficilis.References. Perkins, 1907b: 97 (record: Hawaii, Kilauea); 1910:602 (key); 1913:81

(morphology). Cockerell, 1926:308 (record: Hawaii, Volcano House). Kobayashi, 1974:399(possible pollinator of Argyroxiphium sandwicense; record: Maui, Haleakala). Beardsley,1980:40 (record: Maui, Haleakala).

0.5 mm

A B C D

E F

1.0 mm

Fig. 68. Hylaeus volcanicus, male. A. Seventh sternum, ventral view. B. Eighth sternum, ventralview, and apex, caudal view. C. Eighth sternum, lateral view. D. Genitalia, dorsal view. E. Face withleft scape. F. Scape, lateral view. 0.5 mm scale for A-D; 1 mm scale for E. Extent of pubescenceshown by dashed lines in B-E.


Hylaeus (Nesoprosopis) species A

DIAGNOSIS. Small bees with smoky wings. Female unmarked, with dark, erect hairon T6 and the basal area of the propodeum rugose only on the basal half to third;pronotal collar with strong “shoulders,” lateral portions prominent. Wing length, 4.6mm.

Distribution. Molokai; montane wet forest.Male. Unknown.Female. Head. Malar space medium (ratio about 0.5). Upper paraocular area and

frons with distinct, evenly spaced pits about 1 pit width apart, distorted laterally bylongitudinal striations that curve around the antennal sockets; minutely granular andslightly lustrous from antennal sockets about halfway to median ocellus, dorsal andlateral areas smoothly coriaceous and strongly lustrous; frontal line distinct, forminga distinct groove just above acme. Apex of clypeus weakly emarginate. Dorsal end offacial fovea extended to level of lateral ocellus. Vertex hair 6-10, pale brown, weaklyplumose.

Mesosoma. Fore tarsus with hair apically curved. Lateral area of pronotal collarprominent in dorsal view, almost tuberculate. Scutum with distinct but shallow pits 1-3 pit widths apart, ground unusually coarsely granular, dull; hair mostly rubbed off(the few remaining are 2-6). Scutellum slightly lustrous, with slightly larger pits.Metanotum dull. Mesepisternum with small, shallow, indistinct pits 1-3 pit widthsapart; hair yellowish white, simple, short (long hairs may have been rubbed off); groundminutely granular, lustrous. Hypoepimeron impunctate, lustrous, slightly swollen. Basalarea of propodeum with relatively strong rugae extending a third to halfway to thebrow; remainder including brow smoothly coriaceous.

Metasoma. Disk of T2 impunctate, lightly striate, slightly lustrous, with short, palehairs, about one hair length apart or less. T6 hair long, brown, erect.

Color. Body and legs dark brown to black except pale area on front tibia, apicalyellow mark on front femur, and small obscure orange-brown discolorations at thebases of all tibiae.

Recent collections. Molokai: West Kawela Gulch, 3700 ft.Flower records. Scaevola chamissoniana, Melicope sp. (see remarks).Remarks. A single specimen of this species was collected in West Kawela Gulch

on Molokai. Because it lacks any distinctive features and no male is available, it is notbeing formally described or named; the description above is included to allowidentification of specimens. It is very similar to H. pele, with short body hairs and astrongly convex face; it differs most noticeably in the more sloping dorsal surface ofthe propodeum with a less prominent brow. It is also somewhat larger than that species,which is rarely over 4.0 mm in wing length. Although the specimen was caught flyingaround a flowering Scaevola shrub, it carried a full load of pollen from Melicope, anextremely unusual floral host.


NOMINA DUBIA

The following names are without doubt Ectemnius names mistakenly confused withNesoprosopis. See, for example, Takumi (1999).

“Nesoprosopis rubrocaudatus Blackburn and Cameron,” Suehiro, 1986:41.“Nesoprosopis attripennis Perkins,” Suehiro, 1986:60.


ACKNOWLEDGMENTS

We thank Gordon Nishida of the Bernice P. Bishop Museum, Honolulu (BPBM), who firstencouraged HD to undertake this study and has continued to respond to our many requestsand questions. Loans of additional specimens from BPBM were kindly arranged by G.Allan Samuelson and Keith Arakaki. The manuscript was read critically by Michael S.Engel, James K. Liebherr, and Virginia Scott. Their recommendations were invaluable andgreatly appreciated. We also acknowledge the advice of Roy Snelling. HD’s visits to theHymenoptera Section, The Natural History Museum, London (BMNH), were arrangedwith the kind assistance of George Else, Laraine Ficken, Tom Huddleston, and SuzanneLewis. Likewise, Chris O’Toole welcomed HD to the Hope Entomological Collection,Oxford University Museum of Natural History (OUMNH). The following persons providedspecimens on loan from their institutions (selected collection records are provided in theAppendix): Parker Gambino, formerly Hawaii Volcanoes National Park, Hawaii (HAVO);Bernarr Kumashiro, Department of Agriculture, Plant Pest Control Branch, Honolulu(DOA); Roy Snelling, Los Angeles County Museum, Los Angeles (LACM); Virginia L.Scott, University of Colorado Museum, Boulder (UCM); Arthur C. Medeiros, HaleakalaNational Park, Maui (HALE); Ronald J. McGinley, National Museum of Natural History,Washington (NMNH); Jerry A. Powell, Essig Museum of Entomology, University ofCalifornia at Berkeley (UCB); and Jay A. Rosenheim, University of California at Davis(UCD). Type specimens of the newly described species are deposited in the CornellUniversity Insect Collection (CUIC) and BPBM. We were ably assisted in fieldwork byParker Gambino and Raina Takumi. We also thank Raina Takumi for some of the drawingsof the faces and antennal scapes. For their assistance in facilitating KM’s fieldwork, wewould like to thank David Foote and Linda Pratt of the U.S. Geological Survey, BiologicalResources Division (BRD); Betsy Gagne and Jon Giffin of the Hawaii State Division ofForestry and Wildlife (DOFAW); Tim Tunison, Darcy Hu, and Ron Nagata of the NationalPark Service (NPS); Ed Misaki, Tina Lau, Eric Nishibayashi, Anders Lyons, and JoanYoshioka of The Nature Conservancy; and Randy Bartlett and Hank Oppenheimer of MauiLand & Pineapple Co.

This study was supported in part by a grant (1448-12200-97-G026) from the U. S. Fishand Wildlife Service, Pacific Islands Ecoregion, Ecological Services, Honolulu. We thankAdam Asquith, David R. Hopper, Michael Richardson, Karen W. Rosa, Christa Russell,and Lorena Wada for their attention to the administration of this grant. The field studies ofKM were supported in part from this grant.


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der westpalaarktis (Hymenoptera, Apoidea, Colletidae). Bull. Rech. Agr.Gembloux. 5:745-768

Williams, F. X.1927 Notes on the habits of the bees and wasps of the Hawaiian Islands.

Proc. Hawaii. Entomol. Soc. 6:425-464.Zimmerman, E. C.

1948 Insects of Hawaii. Vol. 1. Introduction. University of Hawai‘i Press,Honolulu.


APPENDIX

Records of Recent Collections

The following are records of recent collections by A. Asquith (AA), H. V. Daly (HD), C.

J. Davis (CD), C. Ewing (CE), D. Foote (DF), C. Fritzler (CF), P. Gambino (PG), J.

Giffin (JG), S. Gon III and U. Kahele (SG), D. Hopper (DH), M. Lopezaraiza (ML), K.

Magnacca (KM), A. C. Medeiros (AM), S. Montgomery (SM), G. Nishida (GN), H.

Oppenheimer (HO), R. P. Papp (RP), D. Polhemus (DP), J. Powell (JP), J. Rosenheim

(JR), V. L. Scott (VS), R. Takumi (RT), and K. R. Wood (KW). Abbreviations for

collection sites are: FR = Forest Reserve; HALE = Haleakala National Park, Maui;

HAVO = Hawaii Volcanoes National Park, Hawaii; KFU = Kona Forest Unit of Hakalau

National Wildlife Refuge, Hawaii; NAR = Natural Area Reserve; NHP = National

Historic Park; PKW = Puu Kukui Watershed; PTA = Pohakuloa Training Area, Hawaii;

SP = State Park, TNC = The Nature Conservancy. Abbreviations for islands are: H =

Hawaii; K = Kauai; Kh = Kahoolawe; L = Lanai; Ma = Maui; Mo = Molokai; O = Oahu.

Species Is. Locality Elev. M F Date Flower/Notes Coll.

akoko H Puu Waawaa 4100 ft 1 1 Aug 2002 Chamaesyce olowaluana KM

andrenoides K Pihea Trail 3 9 Aug 1996 beating Metrosideros, night CE

andrenoides K Alakai Swamp Trail 3600 ft 2 1 21 Aug 1999 Psychotria mariniana KM

andrenoides K Mohihi Trail 3700 ft 1 21 Aug 1999 Chamaesyce atrococca KM

andrenoides K Paaiki Valley 2700 ft 1 1 2 Jul 2000 Claoxylon sandwicense KM

andrenoides K Mohihi Trail 4000 ft 1 8 Jul 2000 Cheirodendron trigynum KM

angustulus Ma Makawao FR 3700 ft 2 1 10 Aug 2002 on Melicope leaves KM

angustulus Ma Makawao FR 4000 ft 3 10 Aug 2002 on Cheirodendron leaves KM

angustulus Ma Makawao FR 4300 ft 1 10 Aug 2002 Rubus penetrans KM

anthracinus H South Point coast 1 17 Jul 1999 Scaevola sericea KM

anthracinus H South Point coast 7 1 18 Jul 1999 Scaevola sericea KM

anthracinus H South Point coast 3 1 Mar 2002 Sida fallax KM

anthracinus H South Point coast 3 1 1 Mar 2002 Tournefortia argentea KM

anthracinus Ma Manawainui west coast 6 1 23 Jun 1999 Sida fallax KM

anthracinus Ma Kanaio NAR 2000 ft 1 23 Jun 1999 Argemone glauca KM

anthracinus Kh Pali O Kalapakea 1000 ft 2 2 14 Jan 2002 Chamaesyce celastroides KW

anthracinus Mo Moomomi Preserve coast 4 1 26 Jun 1999 Scaevola sericea KM

anthracinus Mo Moomomi Preserve coast 1 2 2 Aug 1999 Scaevola sericea KM

anthracinus Mo Moomomi Preserve coast 3 4 Jun 2001 Scaevola sericea KM

anthracinus Mo Moomomi Preserve coast 3 2 4 Jun 2001 over ground KM

anthracinus O Kaena Point NAR coast 1 12 Jun 1999 over ground KM

anthracinus O Kaena Point NAR coast 3 5 12 Jun 1999 Sida fallax KM

anthracinus O Kaena Point NAR coast 1 1 3 Jun 2000 on net KM

anthracinus O Kaena Point NAR coast 3 3 2 Mar 2002 Myoporum sandwicense KM

anthracinus O Kaena Point NAR coast 1 2 Mar 2002 Sida fallax KM

assimulans Ma Lahainaluna 1800 ft 1 1 3 Aug 1999 Sida fallax KM

assimulans Ma Waikapu, Kaohonua 120 m 4 15 Jan 2000 Lipochaeta lobata HO

assimulans Kh Kamohio coast 1 17 Feb 1997 under Prosopis DF

assimulans L Manele Rd. 600 ft 1 17 Jun 1999 Sida fallax KM

assimulans L Polihua Rd. 1000 ft 2 17 Jun 1999 Sida fallax KM

chlorostictus K Kokee Rd. 3600 ft 3 1 24 Aug 1999 Acacia koa KM

chlorostictus K Kokee Rd. 3600 ft 1 24 Aug 1999 Styphelia tameiameiae KM

chlorostictus K Polihale SP coast 8 3 25 Aug 1999 Myoporum sandwicense KM

chlorostictus K Paaiki Valley 2700 ft 1 2 Jul 2000 Claoxylon sandwicense KM



chlorostictus K Nualolo Trail 3200 ft 1 1 3 Jul 2000 over ground KM

chlorostictus K Kukui Trail 2600 ft 2 1 5 Jul 2000 over ground KM

coniceps H Kipuka Maunaiu 1890 m 1 17 Jul 1989 PG

coniceps H 16 mi W Hilo, Hwy 200 4500 ft 3 5 11 Jun 1994 sweeping ferns VS

coniceps H Escape Rd. 3900 ft 1 7 Jan 1999 Metrosideros polymorpha KM

coniceps H Saddle Rd. 5400 ft 3 4 10 Jul 1999 Myoporum sandwicense KM

coniceps H Earthquake Trail 4000 ft 1 21 Jul 1999 Crocosmia crocosmiiflora KM

coniceps H Kipuka Puaulu 4000 ft 3 2 22 Jul 1999 Crocosmia crocosmiiflora KM

coniceps H Earthquake Trail 4000 ft 3 8 Aug 1999 Santalum paniculatum KM

coniceps H Tree Molds 4000 ft 6 3 10 Aug 1999 Santalum paniculatum KM

coniceps H Crater Rim Trail 3800 ft 1 9 Jun 2000 KM

coniceps H Kipuka Puaulu 3900 ft 1 19 Jul 2000 Myoporum sandwicense KM

coniceps H KFU camp 5300 ft 2 1 3 Aug 2000 Myoporum sandwicense KM

coniceps H KFU South Bound. Rd. 4700 ft 1 3 Aug 2000 on Cheirodendron leaves KM

coniceps H KFU South Bound. Rd. 5200 ft 1 3 Aug 2000 Metrosideros polymorpha KM

coniceps H KFU 5500' transect 5600 ft 1 2 3 Aug 2000 Geranium cuneatum KM

coniceps H Puu Huluhulu, N. Hilo 6700 ft 1 27 May 2001 over ground KM

coniceps H Puu Waawaa 5500 ft 1 30 May 2001 Myoporum sandwicense KM

coniceps H HAVO boundary 4000 ft 1 26 Jul 2002 Santalum paniculatum KM

coniceps H Puu Waawaa 4100 ft 4 1 1 Aug 2002 Chamaesyce olowaluana KM

coniceps Ma Haleakala NW slope 2500 m 7 8 24 Jul 1991 malaise trap 6/20-7/24 RT

coniceps Ma Lauulu Trail 2010 m 2 18 Feb 1998 RT

coniceps Ma HALE Service Area 2073 m 2 1 6 Nov 1998 malaise trap 10/30-11/6 RT

coniceps Ma Koolau Gap 6200 ft 1 6 Aug 1999 Metrosideros polymorpha KM

coniceps Ma HALE Service Area 6740 ft 3 20 Mar 2000 malaise trap RT

coniceps Ma Kaupo Trail 6000 ft 1 28 Jun 2000 Scaevola chamissoniana KM

coniceps Ma Puu Keokea 7100 ft 1 20 May 2001 Sophora chrysophylla KM

connectens H Kipuka Nene 2900 ft 1 28 Jun 1998 over ground KM

connectens H Kipuka Nene 2900 ft 1 13 Jan 1999 red pan trap KM

connectens H Kipuka Nene 2900 ft 1 13 Jan 1999 yellow pan trap KM

connectens H Ocean View Estates 4500 ft 1 11 Jul 2000 over ground KM

connectens H Kipuka Nene 2900 ft 1 1 28 Feb 2002 Sophora chrysophylla KM

connectens H Puu Waawaa cone 3820 ft 1 5 Oct 2002 Claoxylon sandwicense JG

connectens H Kipuka Nene 2900 ft 2 28 Feb 2002 Styphelia tameiameiae KM

connectens Ma Waihee Ridge Trail 2000 ft 3 2 5 Aug 1999 Scaevola chamissoniana KM

connectens Ma Puu Kukui Rd. 2600 ft 4 23 Jun 2000 Scaevola chamissoniana KM

connectens Ma Puu Kukui Rd. 2300 ft 1 23 Jun 2000 Metrosideros polymorpha KM

connectens Ma Kahoma 2100 ft 1 22 May 2001 over ground KM

connectens L Munro Trail 3000 ft 1 16 Jun 1999 Metrosideros polymorpha KM

connectens O Puu Kaua 2400 ft 1 11 Jun 1999 Metrosideros polymorpha KM

connectens O Wiliwilinui Trail 2000 ft 2 13 Jun 1999 Metrosideros polymorpha KM

connectens O Wiliwilinui Trail 1800 ft 2 1 26 Jul 1999 Scaevola gaudichaudiana KM

connectens O Puu Kaua 2600 ft 1 1 29 Jul 1999 Scaevola gaudichaudiana KM

connectens O Palikea 3000 ft 1 30 Jul 1999 Broussaisia arguta KM

connectens O Aiea Trail 1700 ft 2 1 2 Jun 2000 Acacia koa KM

connectens O Aiea Trail 1700 ft 2 2 Jun 2000 Metrosideros polymorpha KM

connectens O Lanipo Trail 1500 ft 1 13 Aug 2000 Scaevola gaudichaudiana KM

connectens O Aiea Trail 1700 ft 1 1 14 Aug 2000 Scaevola gaudichaudiana KM

connectens O Aiea Trail 1700 ft 2 1 14 Aug 2000 Styphelia tameiameiae KM

connectens O Aiea Trail 1900 ft 1 14 Aug 2000 Psychotria mariniana KM

connectens O Papali Ridge 1400 ft 1 16 Aug 2000 Scaevola gaudichaudiana KM

connectens O Papali Ridge 1400 ft 1 16 Aug 2000 Metrosideros polymorpha KM

connectens O Papali Ridge 1400 ft 2 1 16 Aug 2000 Chamaesyce multiformis KM

connectens O Pahole NAR 2000 ft 1 8 Jun 2001 Chamaesyce multiformis KM

connectens O Pahole NAR 2100 ft 1 8 Jun 2001 Bidens torta KM



connectens O Pahole NAR 2400 ft 1 18 Feb 2002 Acacia koa KM

connectens O Pahole NAR 2200 ft 2 1 18 Feb 2002 Bidens torta KM

connectens O Pahole NAR 2400 ft 1 5 18 Feb 2002 Canavalia galeata KM

connectens O Pahole NAR 2400 ft 2 1 18 Feb 2002 Rubus rosifolius KM

connectens O Manana Trail 1800 ft 1 19 Feb 2002 Scaevola gaudichaudiana KM

connectens K Kahuamaa Flat 3800 ft 1 24 Aug 1999 Scaevola procera KM

connectens K Polihale SP coast 2 1 25 Aug 1999 Myoporum sandwicense KM

connectens K Paaiki Valley 2700 ft 1 2 Jul 2000 Claoxylon sandwicense KM

connectens K Paaiki Valley 2700 ft 1 3 Jul 2000 Styphelia tameiameiae KM

crabronoides H Olaa Small Tract 3800 ft 1 30 Jul 2000 on ferns KM

crabronoides H KFU South Bound. Rd. 5200 ft 1 2 Aug 2000 Myoporum sandwicense KM

crabronoides H KFU South Bound. Rd. 4700 ft 4 3 Aug 2000 on Cheirodendron leaves KM

crabronoides H KFU South Bound. Rd. 5200 ft 1 3 3 Aug 2000 Myoporum sandwicense KM

crabronoides H Volcano 3600 ft 3 29 Jul 2002 Cheirodendron trigynum KM

difficilis H Kona Coast SP coast 17 18 12 Jun 1994 Tournefortia argentea VS

difficilis H Kipuka Nene 2900 ft 1 4 28 Jun 1998 over ground KM

difficilis H Mauna Loa summit 13677ft 1 14 Jun 1998 over ground KM

difficilis H Kipuka Nene 2900 ft 1 5 Jul 1998 Metrosideros polymorpha KM

difficilis H Mauna Loa Rd. 4200 ft 1 1 2 Jan 1999 over ground KM

difficilis H Kipuka Ki 4200 ft 1 3 Jan 1999 yellow pan trap KM

difficilis H Mauna Loa Rd. 4100 ft 2 3 Jan 1999 yellow pan trap KM

difficilis H Kipuka Nene 2900 ft 9 3 Jan 1999 Sophora chrysophylla KM

difficilis H Kipuka Maunaiu 6000 ft 7 15 4 Jan 1999 over ground KM

difficilis H Kipuka Nene 2900 ft 48 4 Jan 1999 yellow pan trap KM

difficilis H Kipuka Nene 2900 ft 1 4 Jan 1999 red pan trap KM

difficilis H Keanakakoi 3700 ft 1 8 Jan 1999 Metrosideros polymorpha KM

difficilis H Devastation Trail 3700 ft 1 8 Jan 1999 Vaccinium reticulatum KM

difficilis H Makaopuhi Crater 2900 ft 1 1 12 Jan 1999 Metrosideros polymorpha KM

difficilis H Kipuka Nene 2900 ft 4 13 Jan 1999 red pan trap KM

difficilis H KFU South Bound. Rd. 5000 ft 2 8 Jul 1999 Myoporum sandwicense KM

difficilis H KFU South Bound. Rd. 5000 ft 7 3 8 Jul 1999 over ground KM

difficilis H Saddle Rd. 5400 ft 3 4 10 Jul 1999 Myoporum sandwicense KM

difficilis H Kipuka Kalawamana 5000 ft 1 1 13 Jul 1999 Bidens menziesii KM

difficilis H Kipuka Alala 6100 ft 1 13 Jul 1999 over ground KM

difficilis H Kipuka Alala 6100 ft 2 13 Jul 1999 Santalum paniculatum KM

difficilis H Kipuka Alala 5700 ft 3 1 14 Jul 1999 Bidens menziesii KM

difficilis H Kipuka Alala 5800 ft 2 2 14 Jul 1999 Chamaesyce olowaluana KM

difficilis H Kipuka Alala 5800 ft 9 14 Jul 1999 yellow pan trap KM

difficilis H Kaalualu coast 3 3 17 Jul 1999 Tournefortia argentea KM

difficilis H South Point coast 9 7 17 Jul 1999 Scaevola sericea KM

difficilis H Kaalualu coast 4 25 Jul 1999 Myoporum sandwicense KM

difficilis H Lae O Milo coast 4 3 25 Jul 1999 Chamaesyce celastroides KM

difficilis H Kipuka Nene 2900 ft 1 1 13 Aug 1999 over ground KM

difficilis H Kaloko NHP coast 1 14 Aug 1999 Myoporum sandwicense KM

difficilis H Kaloko NHP coast 1 2 15 Aug 1999 Schinus terebinthifolius KM

difficilis H Kaloko NHP coast 5 3 15 Aug 1999 Tournefortia argentea KM

difficilis H Ahumoa 5900 ft 1 7 15 Aug 1999 Bidens menziesii KM

difficilis H Naulu Trail 2900 ft 1 10 11 Mar 2000 over ground DF

difficilis H Kipuka Ki 4200 ft 3 3 6 Jun 2000 Sapindus saponaria KM

difficilis H Kipuka Ki 4200 ft 2 6 Jun 2000 Dodonaea viscosa KM

difficilis H Crater Rim Trail 3800 ft 2 9 Jun 2000 KM

difficilis H Kipuka Ki 4200 ft 2 3 12 Jun 2000 Sapindus saponaria KM

difficilis H Kipuka Ki 4200 ft 1 1 12 Jun 2000 Dodonaea viscosa KM

difficilis H Ocean View Estates 4500 ft 2 11 Jul 2000 over ground KM

difficilis H Ocean View Estates 2400 ft 1 11 Jul 2000 over ground KM



difficilis H Kaloko coast 4 3 13 Jul 2000 Tournefortia argentea KM

difficilis H Hawaii Volcano Obs. 4100 ft 1 24 Jul 2000 Dubautia ciliolata KM

difficilis H Kipuka Nene 2900 ft 1 1 26 Jul 2000 Sophora chrysophylla KM

difficilis H Kipuka Nene 2900 ft 3 1 26 Jul 2000 over ground KM

difficilis H Kipuka Nene 2900 ft 1 1 26 Jul 2000 Styphelia tameiameiae KM

difficilis H Crater Rim Trail 4000 ft 4 28 Jul 2000 Dodonaea viscosa KM

difficilis H Byron’s Ledge 3800 ft 1 29 Jul 2000 Styphelia tameiameiae KM

difficilis H Kilauea Caldera 3600 ft 2 29 Jul 2000 Metrosideros polymorpha KM

difficilis H KFU camp 5300 ft 1 4 2 Aug 2000 Cheirodendron trigynum KM

difficilis H KFU South Bound. Rd. 4700 ft 4 2 3 Aug 2000 on Cheirodendron leaves KM

difficilis H KFU South Bound. Rd. 5200 ft 2 1 3 Aug 2000 Myoporum sandwicense KM

difficilis H Volcano 3750 ft 1 5 Aug 2000 Cheirodendron trigynum KM

difficilis H Volcano 3750 ft 3 6 Aug 2000 Cheirodendron trigynum KM

difficilis H Puu Huluhulu, N. Hilo 6700 ft 3 6 27 May 2001 Myoporum sandwicense KM

difficilis H Puu Huluhulu, N. Hilo 6700 ft 4 3 27 May 2001 over ground KM

difficilis H Ahumoa 5900 ft 2 27 May 2001 over ground KM

difficilis H Saddle Rd. 6200 ft 4 2 27 May 2001 Acacia koa KM

difficilis H Saddle Rd. 6200 ft 1 27 May 2001 Metrosideros polymorpha KM

difficilis H Puu Waawaa 6200 ft 6 30 May 2001 Styphelia tameiameiae KM

difficilis H Puu Waawaa 5500 ft 2 30 May 2001 Myoporum sandwicense KM

difficilis H Kipuka Nene 2900 ft 1 7 28 Feb 2002 Sophora chrysophylla KM

difficilis H South Point coast 21 41 1 Mar 2002 Tournefortia argentea KM

difficilis H Hilina Pali Rd. 3300 ft 2 12 Jul 2002 Dubautia scabra KM

difficilis H Puu Waawaa 4100 ft 3 7 1 Aug 2002 Chamaesyce olowaluana KM

difficilis Ma Kipahulu Valley 1980 m 5 21 May 1998 RT

difficilis Ma Kilohana Pali 2800 m 1 13 Aug 1998 Dubautia menziesii RT

difficilis Ma Sliding Sands Trail 2500 m 2 1 4 Nov 1998 Dubautia menziesii RT

difficilis Ma Kalepeamoa 2835 m 1 14 Dec 1998 RT

difficilis Ma Sliding Sands Trail 9000 ft 1 27 Dec 1998 over ground KM

difficilis Ma Kilohana Pali 2740 m 1 9 Mar 1999 RT

difficilis Ma Kilohana Pali 9400 ft 4 2 22 Jun 1999 over ground KM

difficilis Ma Waikamoi Preserve 6400 ft 3 4 Aug 1999 over ground KM

difficilis Ma Koolau Gap 6200 ft 3 3 6 Aug 1999 Metrosideros polymorpha KM

difficilis Ma above Hosmer Grove 2100 m 1 12 Jun 2000 RT

difficilis Ma PKW fence 2200 ft 4 22 Jun 2000 over ground KM

difficilis Ma Puu Kukui Rd. 2300 ft 3 22 Jun 2000 Metrosideros polymorpha KM

difficilis Ma Puu Kukui Rd. 2300 ft 5 2 23 Jun 2000 Metrosideros polymorpha KM

difficilis Ma Sliding Sands Trail 8650 ft 3 26 Jun 2000 Styphelia tameiameiae KM

difficilis Ma Lauula Trail 6600 ft 1 3 27 Jun 2000 Sophora chrysophylla KM

difficilis Ma Lauula Trail 8000 ft 1 1 27 Jun 2000 over ground KM

difficilis Ma Kaupo Trail 6000 ft 1 2 28 Jun 2000 Metrosideros polymorpha KM

difficilis Ma Kaupo Trail 6000 ft 1 28 Jun 2000 Acacia koa KM

difficilis Ma Kaupo Trail 5500 ft 8 5 28 Jun 2000 over ground KM

difficilis Ma Halemauu Trail 7000 ft 1 29 Jun 2000 Argyroxiphium sandwicense KM

difficilis Ma Laie flats 2040 m 1 26 Jul 2000 RT

difficilis Ma Oili Puu 2050 m 2 27 Jul 2000 RT

difficilis Ma Hanaula, above Puu Anu 915 m 15 3 25 Feb 2000 RT

difficilis Ma Puu Keokea 7100 ft 2 20 May 2001 Styphelia tameiameiae KM

difficilis Ma Puu Keokea 7100 ft 3 20 May 2001 Sophora chrysophylla KM

difficilis Ma Puu Keokea 7100 ft 7 7 20 May 2001 Dodonaea viscosa KM

difficilis Ma Puu Keokea 7100 ft 4 20 May 2001 over ground KM

difficilis Ma Kahoma 2200 ft 1 21 May 2001 over ground KM

difficilis L Lihua 4 19 Aug 1998 SG

difficilis L Munro Trail 3000 ft 1 16 Jun 1999 Metrosideros polymorpha KM

difficilis L Munro Trail 2800 ft 2 7 Aug 1999 Metrosideros polymorpha KM



difficilis Mo Kamakou Rd. 3200 ft 3 3 27 Jun 1999 Metrosideros polymorpha KM

difficilis Mo Puu Kolekole 3400 ft 3 3 28 Jun 1999 over ground KM

difficilis Mo Kamakou Rd. 3900 ft 17 5 2 Jun 2001 Metrosideros polymorpha KM

difficilis Mo Kamakou Rd. 3900 ft 1 3 2 Jun 2001 over ground KM

difficilis Mo Puu Kolekole 3900 ft 3 3 Jun 2001 yellow pan trap KM

dimidiatus H Kipuka Alala 5800 ft 2 14 Jul 1999 Chamaesyce olowaluana KM

dimidiatus H Puu Waawaa 4100 ft 8 5 1 Aug 2002 Chamaesyce olowaluana KM

dumetorum H KFU South Bound. Rd. 5000 ft 2 8 Jul 1999 Myoporum sandwicense KM

dumetorum H Kipuka Puaulu 4000 ft 2 12 Jul 1999 Myoporum sandwicense KM

dumetorum H Kipuka Puaulu 4000 ft 2 3 22 Jul 1999 Crocosmia crocosmiiflora KM

dumetorum H Kipuka Ki 4200 ft 1 12 Jun 2000 Myoporum sandwicense KM

dumetorum H Kipuka Puaulu 3900 ft 1 24 Jul 2000 Myoporum sandwicense KM

dumetorum H Kipuka Puaulu 3900 ft 3 24 Jul 2000 Nestegis sandwicensis KM

dumetorum H Kipuka Puaulu 3900 ft 5 24 Jul 2000 Psychotria hawaiiensis KM

dumetorum H Byron’s Ledge 3800 ft 2 6 29 Jul 2000 on Hedychium leaves KM

dumetorum H Byron’s Ledge 3800 ft 1 29 Jul 2000 Styphelia tameiameiae KM

dumetorum H Byron’s Ledge 3800 ft 2 29 Jul 2000 Myrica faya KM

dumetorum H Byron’s Ledge 3800 ft 1 29 Jul 2000 Fragaria vesca KM

dumetorum H Halemaumau Trail 3700 ft 3 3 29 Jul 2000 Rubus rosifolius KM

dumetorum H Halemaumau Trail 3800 ft 3 29 Jul 2000 Ilex anomala KM

dumetorum H nr. Kilauea Field Station 3900 ft 2 1 30 Jul 2000 Dubautia scabra KM

dumetorum H Olaa Small Tract 3800 ft 1 30 Jul 2000 on ferns KM

dumetorum H KFU South Bound. Rd. 4500 ft 1 2 2 Aug 2000 on Cheirodendron leaves KM

dumetorum H KFU South Bound. Rd. 4700 ft 1 3 Aug 2000 on Cheirodendron leaves KM

dumetorum H KFU South Bound. Rd. 5200 ft 1 3 Aug 2000 Myoporum sandwicense KM

dumetorum H Mountain House Rd. 3000 ft 1 1 25 May 2001 Metrosideros polymorpha KM

dumetorum H Upper Waiakea FR 3600 ft 4 8 26 May 2001 Hedyotis terminalis KM

dumetorum H Upper Hamakua Ditch 3700 ft 1 29 May 2001 Metrosideros polymorpha KM

dumetorum H Olaa Koa Unit 3800 ft 1 27 Feb 2002 Cheirodendron trigynum KM

dumetorum H Kipuka Puaulu 4000 ft 1 31 Jul 2000 Alphitonia ponderosa KM

dumetorum H Kipuka Puaulu 4000 ft 1 31 Jul 2000 Pisonia brunoniana KM

facilis Ma Kokomo 1 25 Oct 1990 PG

filicum H KFU South Bound. Rd. 5200 ft 2 3 Aug 2000 Myoporum sandwicense KM

filicum H Puu Waawaa 4100 ft 2 1 1 Aug 2002 Chamaesyce olowaluana KM

flavifrons K Polihale coast 2 2 17 Jan 1999 Sida fallax KM

flavifrons K Polihale SP coast 5 4 25 Aug 1999 Scaevola sericea KM

flavifrons Ni Lehua, West Horn coast 1 1 19 Feb 2002 Portulaca oleraceae KW

flavipes H Kau, Kaulana Bay coast 1 22 Mar 1993 Sida fallax JP

flavipes H Kau Desert 300 ft 1 9 Aug 1995 Sesbania tomentosa DH

flavipes H Kalu 100 ft 1 22 Aug 1995 Sesbania tomentosa DH

flavipes H Puu Pa 2600 ft 1 7 Nov 1996 SM

flavipes H Kipuka Nene 2900 ft 2 5 Jul 1998 red pan trap KM

flavipes H Kipuka Nene 2900 ft 7 5 Jul 1998 Sesbania tomentosa KM

flavipes H Kipuka Nene 2900 ft 8 5 Jul 1998 red pan trap KM

flavipes H Kipuka Nene 2900 ft 1 3 Jan 1999 over ground KM

flavipes H Hale Pohaku 9000 ft 1 10 Jul 1999 Sophora chrysophylla KM

flavipes H Kipuka Nene 2900 ft 4 11 Jul 1999 over ground KM

flavipes H Kaalualu coast 1 2 17 Jul 1999 Sida fallax KM

flavipes H South Point coast 1 18 Jul 1999 Scaevola sericea KM

flavipes H Kipuka Nene 2900 ft 2 1 26 Jul 2000 Sophora chrysophylla KM

flavipes H South Point coast 2 3 28 May 2001 Jacquemontia ovalifolia KM

flavipes H South Point coast 6 28 May 2001 Sida fallax KM

flavipes H South Point coast 6 40 28 May 2001 Scaevola sericea KM

flavipes H Kipuka Nene 2900 ft 1 28 Feb 2002 Dodonaea viscosa KM

flavipes H Kipuka Nene 2900 ft 3 28 Feb 2002 Styphelia tameiameiae KM



flavipes H Kipuka Nene 2900 ft 1 28 Feb 2002 over ground KM

flavipes H South Point coast 2 9 1 Mar 2002 Tribulus cistoides KM

flavipes H South Point coast 4 1 1 Mar 2002 Jacquemontia ovalifolia KM

flavipes H South Point coast 3 1 Mar 2002 Sida fallax KM

flavipes H South Point coast 3 3 1 Mar 2002 over ground KM

flavipes H South Point coast 2 1 Mar 2002 Tournefortia argentea KM

flavipes L Kahue 1400 ft 6 2 7 Aug 1999 Myoporum sandwicense KM

flavipes L Kahue 1400 ft 3 24 Jun 2000 Myoporum sandwicense KM

flavipes L Kahue 1400 ft 9 7 6 Jun 2001 Myoporum sandwicense KM

flavipes L Kahue 1400 ft 1 6 Jun 2001 Sida fallax KM

fuscipennis Ma Haelaau Cabin 3000 ft 2 3 21 Jun 2000 Metrosideros polymorpha KM

fuscipennis Ma PKW fence 2200 ft 1 22 Jun 2000 Acacia confusa KM

fuscipennis Ma Haelaau Cabin 3000 ft 1 1 23 Jun 2000 on Cheirodendron leaves KM

fuscipennis Ma Puu Kukui Rd. 2300 ft 2 23 Jun 2000 Metrosideros polymorpha KM

fuscipennis Ma Kaulalewelewe 3000 ft 2 10 Aug 2000 Metrosideros polymorpha KM

fuscipennis Ma Kahoma 2600 ft 2 22 May 2001 Metrosideros polymorpha KM

fuscipennis L Munro Trail 3000 ft 2 1 16 Jun 1999 Metrosideros polymorpha KM

fuscipennis L Munro Trail 2800 ft 1 7 Aug 1999 Metrosideros polymorpha KM

fuscipennis L Kaiholena Ridge 2400 ft 1 5 Jun 2001 Cheirodendron trigynum KM

fuscipennis L Munro Trail 2900 ft 1 7 Jun 2001 Tetraplasandra oahuensis KM

fuscipennis L Kaiholena Ridge 2700 ft 1 1 7 Jun 2001 Tetraplasandra oahuensis KM

fuscipennis Mo Puu Kolekole 1200 m 1 16 Jul 1989 JR

fuscipennis Mo Kamakou Rd. 3200 ft 2 2 27 Jun 1999 Metrosideros polymorpha KM

fuscipennis Mo East Kawela Gulch 3600 ft 1 2 Jun 2001 over ferns KM

fuscipennis Mo East Kawela Gulch 3600 ft 1 3 Jun 2001 Psychotria mauiensis KM

haleakalae Ma Puu Kukui Trail 3700 ft 2 3 8 Aug 2000 Cheirodendron trigynum KM

haleakalae Ma Puu Kukui Trail 3700 ft 2 2 10 Aug 2000 Cheirodendron trigynum KM

haleakalae Ma Puu Kukui Trail 3400 ft 2 11 Aug 2000 on Cheirodendron leaves KM

haleakalae Ma Makawao FR 3700 ft 1 10 Aug 2002 on Melicope leaves KM

haleakalae Ma Makawao FR 4300 ft 4 10 Aug 2002 Cheirodendron trigynum KM

haleakalae Mo Kamakou Rd. 3200 ft 1 27 Jun 1999 Metrosideros polymorpha KM

haleakalae Mo West Kawela Gulch 3600 ft 2 2 28 Jun 1999 Scaevola chamissoniana KM

haleakalae Mo Kamakou Rd. 3900 ft 3 2 Jun 2001 Metrosideros polymorpha KM

haleakalae Mo West Kawela Gulch 3700 ft 2 3 2 Jun 2001 Scaevola chamissoniana KM

haleakalae Mo Kawela Rd. 3900 ft 1 1 2 Jun 2001 Cheirodendron trigynum KM

haleakalae Mo East Kawela Gulch 3600 ft 1 2 Jun 2001 over ferns KM

haleakalae Mo Kawela Rd. 3900 ft 1 1 3 Jun 2001 over ferns KM

hilaris Mo Moomomi dunes coast 1 26 Aug 1989 JR

hilaris Mo Moomomi Preserve coast 1 30 Jun 1999 over ground KM

hirsutulus K Alakai Wilderness 1 1 2 Nov 1999 Metrosideros DH

hirsutulus K Nualolo Cliff Trail 2800 ft 5 3 Jul 2000 Styphelia tameiameiae KM

hirsutulus K Milolii Rd. 2700 ft 1 7 Jul 2000 over ground KM

hirsutulus K Mohihi Trail 3900 ft 1 8 Jul 2000 Chamaesyce atrococca KM

hostilis K Polihale SP coast 4 25 Aug 1999 Scaevola sericea KM

hostilis K Awaawapuhi Trail 3400 ft 1 3 Jul 2000 over ground KM

hostilis K Nualolo Cliff Trail 2800 ft 1 1 3 Jul 2000 Styphelia tameiameiae KM

hostilis K Awaawapuhi Trail 3400 ft 1 1 4 Jul 2000 over ground KM

hula H Tree Molds 4000 ft 5 10 Aug 1999 Santalum paniculatum KM

hula H Tree Molds 4000 ft 1 10 Jul 2002 Santalum paniculatum KM

hula H HAVO boundary fence 4000 ft 1 26 Jul 2002 Santalum paniculatum KM

hula H Puu Waawaa 4100 ft 2 1 1 Aug 2002 Chamaesyce olowaluana KM

inquilina H south of Bird Park 1230 m 1 6 4 Aug 1989 PG

inquilina H Kipuka Alala 5800 ft 1 9 Mar 1992 Myoporum sandwicense GN

inquilina H Mauna Loa Rd. 2050 m 2 1 21 Mar 1993 JP

inquilina H 0.9 mi. S Mauna Loa Rd. 6000 ft 2 1 4 Jan 1999 over ground KM



inquilina H Kipuka Nene 2900 ft 1 13 Jan 1999 over ground KM

inquilina H near Tree Molds 4000 ft 1 11 Jun 2000 over ground KM

kauaiensis K Mt. Kahili 2500 ft 3 9 1 Oct 1997 malaise trap Aug.-Oct. AA

kauaiensis K Alakai Swamp Trail 3700 ft 2 3 Jul 1999 Metrosideros polymorpha KM

kauaiensis K Alakai Swamp Trail 3700 ft 2 3 Jul 1999 Scaevola procera KM

kauaiensis K Na Pali-Kona FR 4100 ft 2 1 4 Jul 1999 Scaevola procera KM

kauaiensis K Alakai Swamp Trail 3700 ft 1 5 Jul 1999 Metrosideros polymorpha KM

kauaiensis K Alakai Swamp Trail 3600 ft 1 21 Aug 1999 Scaevola procera KM

kauaiensis K Alakai Swamp Trail 3600 ft 1 2 21 Aug 1999 Metrosideros polymorpha KM

kauaiensis K Kahuamaa Flat 3800 ft 1 24 Aug 1999 Styphelia tameiameiae KM

kauaiensis K Mohihi Trail 4000 ft 2 8 Jul 2000 Metrosideros polymorpha KM

kokeensis K Kokee Rd. 3600 ft 4 24 Aug 1999 Acacia koa KM

kokeensis K Kokee Rd. 3600 ft 1 24 Aug 1999 yellow sticky card KM

kokeensis K Awaawapuhi Trail 3400 ft 11 3 Jul 2000 Chamaesyce atrococca KM

kokeensis K Awaawapuhi Trail 3400 ft 8 12 4 Jul 2000 Chamaesyce atrococca KM

kokeensis K Awaawapuhi Trail 3400 ft 1 8 Jul 2000 Chamaesyce atrococca KM

kona H Kipuka Alala 5700 ft 1 14 Jul 1999 Bidens menziesii KM

kona H Kipuka Alala 5800 ft 9 14 Jul 1999 Chamaesyce olowaluana KM

kuakea O Moho Gulch Ridge 1900 ft 2 1 Aug 1997 over ground DH

kukui Ma Puu Kukui Trail 3000 ft 6 1 11 Aug 2000 over ground KM

laetus H Kipuka Nene 2900 ft 1 28 Jun 1998 over ground KM

laetus H Kipuka Nene 2900 ft 1 5 Jul 1998 Metrosideros polymorpha KM

laetus H Kipuka Nene 2900 ft 1 3 Jan 1999 over ground KM

laetus H Kipuka Nene 2900 ft 1 2 4 Jan 1999 yellow pan trap KM

laetus H Kipuka Nene 2900 ft 1 1 13 Jan 1999 yellow pan trap KM

laetus H Kipuka Nene 2900 ft 1 13 Jan 1999 yellow pan trap KM

laetus H Kipuka Nene 2900 ft 5 11 Jul 1999 over ground KM

laetus H Kipuka Kalawamana 5000 ft 6 7 13 Jul 1999 Bidens menziesii KM

laetus H Kipuka Alala 6100 ft 1 13 Jul 1999 over ground KM

laetus H Kipuka Alala 5800 ft 1 1 14 Jul 1999 Chamaesyce olowaluana KM

laetus H Kipuka Alala 5400 ft 2 6 14 Jul 1999 yellow pan trap KM

laetus H Ahumoa 5900 ft 3 15 Aug 1999 Bidens menziesii KM



laetus H Kipuka Nene 2900 ft 1 2 26 Jul 2000 Styphelia tameiameiae KM

laetus H Kipuka Nene 2900 ft 8 28 Feb 2002 Dodonaea viscosa KM

laetus H Kipuka Nene 2900 ft 1 28 Feb 2002 Sophora chrysophylla KM

laetus H Kipuka Nene 2900 ft 1 4 28 Feb 2002 Styphelia tameiameiae KM

laetus H Hilina Pali Rd. 2000 ft 6 12 Jul 2002 Plantago lanceolata KM

laetus H Hilina Pali Rd. 2500 ft 3 1 12 Jul 2002 Myoporum sandwicense KM

laetus H Hilina Pali Rd. 2500 ft 3 1 12 Jul 2002 Metrosideros polymorpha KM

laetus H HAVO boundary fence 4000 ft 1 26 Jul 2002 Santalum paniculatum KM

laetus H Puu Waawaa 4100 ft 3 4 1 Aug 2002 Chamaesyce olowaluana KM

laetus H Humuula Saddle 6800 ft 1 1 Aug 2002 Chamaesyce olowaluana KM

laetus Ma Lahainaluna 1800 ft 3 5 3 Aug 1999 Santalum freycinetianum KM

laetus Ma Lahainaluna 1800 ft 5 3 Aug 1999 Sida fallax KM

laetus Ma Kahoma 2100 ft 1 23 May 2001 over ground KM

laetus L Lihua 4 19 Aug 1998 SG

laetus L nr. Puu Maua 1 19 Aug 1998 SG

laetus L Kanepuu Pres., Kahue 1600 ft 7 6 15 Jun 1999 Sida fallax KM

laetus L Garden of the Gods 1400 ft 3 17 Jun 1999 over ground KM

laetus L Kahue 1400 ft 12 9 7 Aug 1999 Myoporum sandwicense KM

laetus L Kahue 1400 ft 1 24 Jun 2000 Myoporum sandwicense KM

laetus L Kahue 1400 ft 10 1 6 Jun 2001 Myoporum sandwicense KM

laetus L Kahue 1400 ft 1 1 6 Jun 2001 Sida fallax KM



laetus L Kaiholena Ridge 2600 ft 1 7 Jun 2001 Metrosideros polymorpha KM

laetus O Pahole NAR 2000 ft 1 8 Jun 2001 Chamaesyce multiformis KM

laetus K Black Pipe Trail 3500 ft 1 2 Jul 1999 Styphelia tameiameiae KM

laetus K Iliau Loop 3000 ft 10 2 5 Jul 1999 Bidens sandvicensis KM

laetus K Alakai Swamp Trail 3600 ft 1 1 21 Aug 1999 Metrosideros polymorpha KM

laetus K Alakai Swamp Trail 3600 ft 1 21 Aug 1999 over ground KM

laetus K Kokee Rd. 3600 ft 1 24 Aug 1999 Acacia koa KM

laetus K Kokee Rd. 3600 ft 4 1 24 Aug 1999 Styphelia tameiameiae KM

laetus K Milolii Rd. 2600 ft 1 24 Aug 1999 Styphelia tameiameiae KM

laetus K Paaiki Valley 2700 ft 2 1 2 Jul 2000 Claoxylon sandwicense KM

laetus K Milolii Rd. 3050 ft 3 2 Jul 2000 Acacia koa KM

laetus K Awaawapuhi Trail 3400 ft 2 1 3 Jul 2000 Chamaesyce atrococca KM

laetus K Nualolo Cliff Trail 2800 ft 2 1 3 Jul 2000 Styphelia tameiameiae KM

laetus K Mohihi Trail 3900 ft 3 2 8 Jul 2000 Chamaesyce atrococca KM

longiceps Ma Waiehu dune 100 ft 4 2 5 Aug 1999 Scaevola coriacea KM

longiceps Ma Waiehu dune 100 ft 1 21 May 2001 over ground KM

longiceps L Polihua Rd. 1000 ft 1 1 15 Jun 1999 Sida fallax KM

longiceps L Polihua Rd. 1000 ft 5 5 17 Jun 1999 Sida fallax KM

longiceps L Kahue 1400 ft 1 2 7 Aug 1999 Sida fallax KM

longiceps L Shipwreck Beach coast 2 6 Jun 2001 Scaevola sericea KM

longiceps Mo Moomomi Preserve coast 2 1 26 Jun 1999 Scaevola sericea KM

longiceps Mo Moomomi Preserve coast 1 2 30 Jun 1999 Scaevola sericea KM

longiceps Mo Moomomi Preserve coast 1 1 2 Aug 1999 Scaevola sericea KM

longiceps O Kaena Point NAR coast 3 6 12 Jun 1999 Sesbania tomentosa KM

longiceps O Kaena Point NAR coast 9 4 12 Jun 1999 Scaevola sericea KM

longiceps O Kaena Point NAR coast 2 2 3 Jun 2000 Sesbania tomentosa KM

longiceps O Kaena Point NAR coast 3 3 Jun 2000 on net KM

longiceps O Kaena Point NAR coast 6 17 2 Mar 2002 Sesbania tomentosa KM

longiceps O Kaena Point NAR coast 4 12 2 Mar 2002 Myoporum sandwicense KM

longiceps O Kaena Point NAR coast 2 5 2 Mar 2002 Sida fallax KM

longiceps O Kaena Point NAR coast 1 2 Mar 2002 Santalum ellipticum KM

longiceps O Kaena Point NAR coast 1 2 2 Mar 2002 Chamaesyce degeneri KM

longiceps O Kaena Point NAR coast 1 2 Mar 2002 Vitex rotundifolia KM

mana O Manana Trail 1400 ft 3 1 3 Mar 2002 Santalum freycinetianum KM

mimicus O Wiliwilinui Trail 2000 ft 1 13 Jun 1999 Metrosideros polymorpha KM

mimicus O Wiliwilinui Trail 1800 ft 2 1 26 Jul 1999 Scaevola gaudichaudiana KM

mimicus O Aiea Trail 1700 ft 3 2 Jun 2000 Acacia koa KM

mimicus O Aiea Trail 1700 ft 3 14 Aug 2000 Scaevola gaudichaudiana KM

mimicus O Aiea Trail 1900 ft 1 14 Aug 2000 Ilex anomala KM

mimicus O Aiea Trail 1900 ft 1 14 Aug 2000 Metrosideros polymorpha KM

mimicus O Aiea Trail 1900 ft 4 1 14 Aug 2000 Psychotria mariniana KM

muranus H Volcano 3750 ft 4 5 Aug 2000 Cheirodendron trigynum KM

muranus H Volcano 3750 ft 10 6 Aug 2000 Cheirodendron trigynum KM

muranus H Volcano 3750 ft 2 29 Jul 2002 Cheirodendron trigynum KM

muranus H Puu Puai 3700 ft 1 10 Jul 2002 Metrosideros polymorpha ML

mutatus K Kahuamaa Flat 3800 ft 4 24 Aug 1999 Styphelia tameiameiae KM

mutatus K Makaha Rd. 3500 ft 1 24 Aug 1999 Styphelia tameiameiae KM

mutatus K Paaiki Valley 2700 ft 3 1 2 Jul 2000 Claoxylon sandwicense KM

mutatus K Awaawapuhi Trail 3400 ft 1 3 Jul 2000 Chamaesyce atrococca KM

mutatus K Awaawapuhi Trail 3400 ft 6 3 4 Jul 2000 Chamaesyce atrococca KM

mutatus K Kuia Valley 2400 ft 1 7 Jul 2000 Antidesma platyphyllum KM

nivicola Ma Haleakala W slope 8400 ft 1 29 Jul 1984 Styphelia tameiameiae AM

nivicola Ma Haleakala W slope 6800 ft 1 8 May 1985 in silk tube nest under rock AM

nivicola Ma Holua Cabin 7200 ft 1 6 Jun 1985 Oenothera CF

nivicola Ma Haleakala Crater 7600 ft 1 25 Aug 1986 Argyroxiphium sandwicense AM



nivicola Ma Haleakala NW slope 2080 m 5 22 May 1991 PG

nivicola Ma below Halemauu Trail 2300 m 4 20 Jun 1991 RT

nivicola Ma above Halemauu Trail 2600 m 1 2 20 Jun 1991 RT

nivicola Ma Haleakala NW slope 2500 m 35 51 24 Jul 1991 malaise trap 6/14-7/24 RT

nivicola Ma Halemauu Trail 1 5 27 May 1992 RT

nivicola Ma Halemauu Trail 7840 ft 4 14 7 Sep 1993 basking and on flowers HD

nivicola Ma Halemauu Trail 7840 ft 1 7 Sep 1993 Geranium cuneatum HD

nivicola Ma Holua Trail 8000 ft 1 18 Jun 1995 DH

nivicola Ma base of Puu Maile 2195 m 1 25 Jun 1996 RT

nivicola Ma Sliding Sands Trail 2590 m 1 25 Jun 1996 RT

nivicola Ma Paliku 1830 m 1 26 Jun 1996 RT

nivicola Ma Mamane Hill 2255 m 1 28 Jun 1996 Styphelia tameiameiae RT

nivicola Ma Halemauu Trail 2440 m 1 28 Aug 1996 RT

nivicola Ma near HALE visitor center 9750 ft 1 11 Sep 1996 Dubautia menziesii RT

nivicola Ma near Kilohana Peak 2880 m 1 12 Sep 1996 RT

nivicola Ma Mees Solar Observ. 3050 m 1 16 Sep 1996 Dubautia menziesii RT

nivicola Ma above Halemauu Trail 1 25 Sep 1996 RT

nivicola Ma Sliding Sands Trail 2500 m 1 27 Oct 1996 RT

nivicola Ma near Kapalaoa Cabin 2195 m 2 11 Feb 1997 RT

nivicola Ma Silversword Loop 2195 m 1 2 14 Feb 1997 Styphelia tameiameiae RT

nivicola Ma Halemauu switchbacks 2195 m 1 1 3 Mar 1997 RT

nivicola Ma Halemauu Trail 2250 m 4 3 3 Mar 1997 RT

nivicola Ma Puu Maile 2195 m 1 10 Mar 1997 RT

nivicola Ma Kalahaku 2835 m 2 14 Mar 1997 Styphelia tameiameiae RT

nivicola Ma Kilohana Peak 2890 m 3 1 14 Mar 1997 RT

nivicola Ma below Kilohana Peak 2500 m 5 9 4 Apr 1997 Styphelia tameiameiae RT

nivicola Ma Laie 2040 m 1 1 7 Apr 1997 RT

nivicola Ma Lauulu Trail 2440 m 5 4 15 May 1997 RT

nivicola Ma Kaluaiki 2195 m 2 27 May 1997 RT

nivicola Ma Lauulu Trail 2465 m 5 6 27 May 1997 RT

nivicola Ma Haleakala W slope 2900 m 4 1 13 Jun 1997 RT

nivicola Ma HALE offices 2073 m 1 3 17 Jun 1997 RT

nivicola Ma near White Hill 2860 m 9 15 Jul 1997 Argyroxiphium sandwicense RT

nivicola Ma Kuiki 2170 m 1 3 7 Aug 1997 RT

nivicola Ma Haleakala NW slope 2310 m 1 12 Aug 1997 RT

nivicola Ma Science City 3010 m 1 13 Aug 1997 RT

nivicola Ma Laie flats 2060 m 2 19 Aug 1997 RT

nivicola Ma Na Mana o ke Akua 2194 m 2 1 21 Aug 1997 RT

nivicola Ma Paliku 1950 m 1 1 25 Sep 1997 RT

nivicola Ma near Mees Obs. 3010 m 1 28 Oct 1997 RT

nivicola Ma wash below Kapalaoa 2165 m 1 1 13 Jan 1998 RT

nivicola Ma Lauulu Trail 2015 m 1 14 Jan 1998 RT

nivicola Ma Kalepeamoa 2835 m 1 14 Dec 1998 Styphelia tameiameiae RT

nivicola Ma Sliding Sands Trail 9000 ft 1 1 27 Dec 1998 over ground KM

nivicola Ma Leleiwi 2710 m 1 16 Apr 1999 Styphelia tameiameiae RT

nivicola Ma Halemauu Trail 8000 ft 3 2 12 May 1999 over ground RT

nivicola Ma Haleakala NW slope 2250 m 1 12 May 1999 RT

nivicola Ma Halemauu Trail 8000 ft 5 2 19 Jun 1999 over ground KM

nivicola Ma Supply Trail 6800 ft 16 25 Jun 1999 yellow pan trap KM

nivicola Ma HALE HQ 7200 ft 1 1 3 Aug 1999 Argyroxiphium sandwicense KM

nivicola Ma HALE Service Area 2073 m 10 4 29 Feb 2000 malaise trap 2/24-2/29 RT

nivicola Ma Halemauu Trail 7900 ft 2 18 Jun 2000 Styphelia tameiameiae KM

nivicola Ma Halemauu Trail 7900 ft 2 18 Jun 2000 over ground KM

nivicola Ma Halemauu Trail 7900 ft 1 25 Jun 2000 over ground KM

nivicola Ma Lauula Trail 8000 ft 4 1 27 Jun 2000 over ground KM



nivicola Ma Puu Keokea 7100 ft 6 8 20 May 2001 Styphelia tameiameiae KM

nivicola Ma Puu Keokea 7100 ft 2 7 20 May 2001 Sophora chrysophylla KM

nivicola Ma Puu Keokea 7100 ft 1 20 May 2001 Dodonaea viscosa KM

ombrias H Kipuka Kalawamana 5000 ft 10 3 13 Jul 1999 Bidens menziesii KM

ombrias H South Point coast 3 3 17 Jul 1999 Sida fallax KM

ombrias H South Point coast 1 18 Jul 1999 Scaevola sericea KM

ombrias H Ahumoa 5900 ft 1 15 Aug 1999 Bidens menziesii KM

ombrias H South Point coast 1 28 May 2001 Scaevola sericea KM

ombrias H South Point coast 3 1 Mar 2002 Tribulus cistoides KM

ombrias H South Point coast 1 1 Mar 2002 Sida fallax KM

ombrias H Puu Waawaa 4300 ft 1 26 Sep 2002 Chamaesyce olowaluana JG

paradoxicus H Kipuka Alala 5800 ft 5 1 14 Jul 1999 Chamaesyce olowaluana KM

paradoxicus H Puu Waawaa 4100 ft 2 1 Aug 2002 Chamaesyce olowaluana KM

paradoxicus H Puu Waawaa 4100 ft 1 1 Aug 2002 Myoporum sandwicense KM

pele H Mauna Loa Rd. 4100 ft 2 1 2 Jan 1999 Sophora chrysophylla KM

pele H Kipuka Nene 2900 ft 1 4 Jan 1999 yellow pan trap KM

pele H Kipuka Kalawamana 5000 ft 1 13 Jul 1999 Bidens menziesii KM

pele H Kipuka Alala 5700 ft 5 2 14 Jul 1999 Bidens menziesii KM

pele H Kipuka Alala 5800 ft 5 1 14 Jul 1999 Chamaesyce olowaluana KM

pele H Kipuka Puaulu 4000 ft 2 22 Jul 1999 Crocosmia crocosmiiflora KM

pele H Ahumoa 5900 ft 1 3 15 Aug 1999 Bidens menziesii KM

pele H Kipuka Puaulu 3900 ft 5 19 Jul 2000 Myoporum sandwicense KM

pele H Kipuka Puaulu 3900 ft 1 24 Jul 2000 Myoporum sandwicense KM

pele H Kipuka Nene 2900 ft 1 2 26 Jul 2000 Sophora chrysophylla KM

pele H Kipuka Nene 2900 ft 2 26 Jul 2000 over ground KM

pele H Puu Huluhulu, N. Hilo 6700 ft 1 27 May 2001 Myoporum sandwicense KM

pele H Kipuka Nene 2900 ft 1 7 28 Feb 2002 Sophora chrysophylla KM

pele H Puu Waawaa 4100 ft 3 1 Aug 2002 Chamaesyce olowaluana KM

psammobius Ma Eleilei Bay coast 7 6 8 Aug 2002 Sesuvium portulacastrum KM

psammobius Ma Eleilei Bay coast 3 1 8 Aug 2002 Bacopa monnieri KM

pubescens H Kau Desert trailhead 930 m 1 6 Dec 1988 PG

pubescens H Kilauea Iki rim 966 m 4 1 13 Apr 1989 PG

pubescens H Kilauea Iki rim 966 m 3 25 Apr 1989 PG

pubescens H Thurston Lava Tube 1 9 Jun 1989 PG

pubescens H Bird Park 1230 m 1 5 Jul 1989 PG

pubescens H Bird Park 1230 m 3 25 Jul 1989 PG

pubescens H Olaa Tract 1 21 Oct 1989 PG

pubescens H Kilauea Field Station 1175 m 1 2 Oct 1990 PG

pubescens H Lua Manu crater 3700 ft 1 1 1 Jan 1999 Metrosideros polymorpha KM

pubescens H Kilauea Field Station 3800 ft 1 1 Jan 1999 nest in Metrosideros KM

pubescens H Tree Planting Rd. 4200 ft 1 5 Jan 1999 Metrosideros polymorpha KM

pubescens H Devastation Trail 3700 ft 3 2 8 Jan 1999 Metrosideros polymorpha KM

pubescens H Hilina Pali Rd. 3300 ft 1 11 Jan 1999 red pan trap KM

pubescens H Kilauea Iki 3900 ft 1 5 Jun 1999 over ground KM

pubescens H Tree Molds 4000 ft 1 9 Aug 1999 Santalum paniculatum KM

pubescens H Tree Molds 4000 ft 1 10 Aug 1999 Santalum paniculatum KM

pubescens H Olaa Small Tract 3800 ft 1 15 Jun 2000 Metrosideros polymorpha KM

pubescens H Kipuka Puaulu 3900 ft 1 24 Jul 2000 Nestegis sandwicensis KM

pubescens H Kilauea Caldera 3600 ft 1 29 Jul 2000 Metrosideros polymorpha KM

pubescens H Olaa, Wright Rd. 3800 ft 1 30 Jul 2000 Cheirodendron trigynum KM

pubescens H KFU camp 5300 ft 1 2 Aug 2000 Cheirodendron trigynum KM

pubescens H Volcano 3750 ft 2 6 Aug 2000 Cheirodendron trigynum KM

pubescens H Mountain House Rd. 3000 ft 1 2 25 May 2001 Metrosideros polymorpha KM

pubescens H Mountain House Rd. 3000 ft 1 25 May 2001 Tibouchina urvilleana KM

pubescens H Upper Hamakua Ditch 3700 ft 1 29 May 2001 Metrosideros polymorpha KM



pubescens H Olaa Koa Unit 3800 ft 3 27 Feb 2002 over ferns KM

pubescens H Kipuka Puaulu 4000 ft 1 31 Jul 2002 Psychotria hawaiiensis KM

pubescens H South Kona FR 4700 ft 1 4 Aug 2002 Myoporum sandwicense KM

rugulosus H below 3-Tree Kipuka 1890 m 1 9 Jul 1989 Metrosideros polymorpha PG

rugulosus H Kipuka Nene 2900 ft 1 16 May 1998 yellow pan trap KM

rugulosus H Kilauea Military Camp 4000 ft 1 12 Jul 2002 Metrosideros polymorpha ML

rugulosus H Napau Trail 2900 ft 5 9 14 Jul 2002 around stones KM

rugulosus H Puu Puai 3700 ft 1 26 Jul 2002 Metrosideros polymorpha ML

setosifrons H Earthquake Trail 4000 ft 1 9 Aug 1999 Santalum paniculatum KM

setosifrons H Tree Molds 4000 ft 1 9 Aug 1999 Santalum paniculatum KM

setosifrons H Tree Molds 4000 ft 1 10 Aug 1999 Santalum paniculatum KM

setosifrons H Mauna Loa Est. 3600 ft 1 20 May 2000 DF

setosifrons H Volcano 3750 ft 1 5 Aug 2000 Cheirodendron trigynum KM

setosifrons H Kipuka Puaulu 4000 ft 1 31 Jul 2002 Psychotria hawaiiensis KM

solaris K Keoneloa Bay coast 3 2 19 Aug 1989 JP

solaris K Polihale SP coast 1 2 Feb 1994 Sida fallax DH

solaris K Polihale SP coast 10 11 25 Aug 1999 Scaevola sericea KM

specularis H KFU South Bound. Rd. 5000 ft 1 8 Jul 1999 Myoporum sandwicense KM

specularis H KFU South Bound. Rd. 5200 ft 6 2 Aug 2000 Myoporum sandwicense KM

specularis H KFU camp 5300 ft 1 3 Aug 2000 Myoporum sandwicense KM

specularis H KFU South Bound. Rd. 5200 ft 8 4 3 Aug 2000 Myoporum sandwicense KM

specularis O Manana Trail 1600 ft 1 19 Feb 2002 Metrosideros polymorpha KM

specularis K Na Pali-Kona FR 4100 ft 1 4 Jul 1999 Scaevola procera KM

specularis K Awaawapuhi Trail 3800 ft 3 1 4 Jul 2000 Claoxylon sandwicense KM

specularis K Awaawapuhi Trail 3800 ft 3 5 Jul 2000 Claoxylon sandwicense KM

sphecodoides H Kipuka Nene 2900 ft 6 11 Jul 1999 over ground KM

sphecodoides H Kipuka Alala 6100 ft 1 13 Jul 1999 over ground KM

sphecodoides H Kipuka Nene 2900 ft 1 26 Jul 2000 over ground KM

sphecodoides H South Point coast 1 1 Mar 2002 over ground KM

sphecodoides H Keauhou Ranch 4500 ft 2 29 Jun 2002 Cheirodendron trigynum KM

takumiae Ma Haleakala NW slope 2500 m 7 36 24 Jul 1991 malaise trap 6/14-7/24 RT

takumiae Ma Haleakala W slope 2865 m 1 13 Jun 1997 RT

unicus Ma Kipahulu Valley 945 m 1 7 Jan 1997 RT

unicus Ma Puu Kukui Rd. 2300 ft 3 22 Jun 2000 Metrosideros polymorpha KM

unicus Ma Haelaau Cabin 3000 ft 1 23 Jun 2000 Cheirodendron trigynum KM

unicus Ma Puu Kukui Rd. 2300 ft 3 1 23 Jun 2000 Metrosideros polymorpha KM

unicus Ma Puu Kukui Trail 3700 ft 2 2 8 Aug 2000 Cheirodendron trigynum KM

unicus Ma Puu Kukui Trail 3700 ft 5 5 10 Aug 2000 Cheirodendron trigynum KM

unicus Ma Puu Kukui Trail 3200 ft 1 11 Aug 2000 Dubautia laxa KM

unicus Ma Makawao FR 4300 ft 2 10 Aug 2002 Cheirodendron trigynum KM

unicus L Munro Trail 3000 ft 4 5 16 Jun 1999 Metrosideros polymorpha KM

unicus L Munro Trail 3000 ft 1 16 Jun 1999 Scaevola chamissoniana KM

unicus L Munro Trail 2900 ft 1 7 Jun 2001 Tetraplasandra oahuensis KM

unicus Mo Kamakou Rd. 3200 ft 1 27 Jun 1999 Metrosideros polymorpha KM

unicus Mo West Kawela Gulch 3600 ft 2 28 Jun 1999 Scaevola chamissoniana KM

unicus Mo Kamakou Rd. 3900 ft 1 2 Jun 2001 Metrosideros polymorpha KM

unicus Mo Pepeopae boardwalk 4000 ft 1 1 2 Jun 2001 over ferns KM

unicus O Mt. Kaala 4000 ft 2 1 Oct 1997 nest in Bidens torta twig DH

unicus O Wiliwilinui Trail 2000 ft 3 3 13 Jun 1999 Metrosideros polymorpha KM

unicus O Wiliwilinui Trail 1800 ft 1 1 26 Jul 1999 Scaevola gaudichaudiana KM

unicus O Puu Kaua 2600 ft 3 1 29 Jul 1999 Scaevola gaudichaudiana KM

unicus O Koolau Summit Trail 2300 ft 1 1 Jun 2000 Psychotria fauriei KM

unicus O Papali Ridge 1500 ft 2 4 Jun 2000 Metrosideros polymorpha KM

unicus O Aiea Trail 1900 ft 2 14 Aug 2000 Metrosideros polymorpha KM

unicus O Aiea Trail 1900 ft 1 14 Aug 2000 Tetraplasandra oahuensis KM



unicus O Papali Ridge 1400 ft 1 16 Aug 2000 Metrosideros polymorpha KM

unicus O Manana Trail 1600 ft 4 19 Feb 2002 Metrosideros polymorpha KM

unicus O Manana Trail 1800 ft 1 2 19 Feb 2002 Acacia koa KM

unicus O Manana Trail 1800 ft 1 19 Feb 2002 Scaevola gaudichaudiana KM

unicus O Manana Trail 1600 ft 1 3 Mar 2002 Metrosideros polymorpha KM

volatilis Ma Haleakala NW slope 2080 m 1 22 May 1991 PG

volatilis Ma below Halemauu Trail 2300 m 1 20 Jun 1991 RT

volatilis Ma above Halemauu Trail 2600 m 1 20 Jun 1991 RT

volatilis Ma Haleakala NW slope 2500 m 9 13 24 Jul 1991 malaise trap 6/14-7/24 RT

volatilis Ma Haleakala N slope 2150 m 1 13 Aug 1991 PG

volatilis Ma Halemauu Trail 1 2 27 May 1992 RT

volatilis Ma near Halemauu Trail 7840 ft 1 7 Sep 1993 HD

volatilis Ma Paliku 1830 m 1 26 Jun 1996 RT

volatilis Ma near Halemauu Trail 2375 m 1 1 Oct 1996 RT

volatilis Ma Puu Maile 2195 m 1 10 Mar 1997 RT

volatilis Ma Kalahaku 2835 m 1 14 Mar 1997 Styphelia tameiameiae RT

volatilis Ma Laie 2040 m 3 7 Apr 1997 RT

volatilis Ma Waikane 2010 m 2 8 Apr 1997 RT

volatilis Ma Kaluaiki 2225 m 1 1 15 May 1997 RT

volatilis Ma Lauulu Trail 1955 m 1 2 27 May 1997 RT

volatilis Ma Lauulu Trail 2465 m 2 1 27 May 1997 RT

volatilis Ma Kuiki 2170 m 4 3 7 Aug 1997 RT

volatilis Ma Paliku 1950 m 1 25 Sep 1997 RT

volatilis Ma Oili Puu 2040 m 1 6 Aug 1998 RT

volatilis Ma Halemauu Trail 8000 ft 2 1 12 May 1999 over ground RT

volatilis Ma Haleakala NW slope 2250 m 1 12 May 1999 RT

volatilis Ma Halemauu Trail 8000 ft 2 6 19 Jun 1999 over ground KM

volatilis Ma Halemauu Trail 7800 ft 8 4 23 Jun 1999 over ground KM

volatilis Ma Supply Trail 6800 ft 1 1 25 Jun 1999 yellow pan trap KM

volatilis Ma Lahainaluna 1800 ft 1 3 Aug 1999 over ground KM

volatilis Ma Waikamoi Preserve 6400 ft 1 4 Aug 1999 over ground KM

volatilis Ma below Halemauu Trail 2370 m 1 5 Aug 1999 RT

volatilis Ma Lauulu Trail 2130 m 2 9 Sep 1999 RT

volatilis Ma HALE Service Area 6800 ft 1 20 Mar 2000 malaise trap RT

volatilis Ma HALE visitor center 2975 m 1 28 May 2000 RT

volatilis Ma Laie flats 2040 m 2 31 May 2000 RT

volatilis Ma above Hosmer Grove 2100 m 1 12 Jun 2000 RT

volatilis Ma near Koolau 2250 m 3 1 12 Jun 2000 RT

volatilis Ma Halemauu Trail 7900 ft 1 25 Jun 2000 over ground KM

volatilis Ma Lauula Trail 8000 ft 1 27 Jun 2000 over ground KM

volatilis Ma Kalapawili Ridge 7400 ft 1 1 27 Jun 2000 over ground KM

volatilis Ma Kaupo Trail 5500 ft 1 2 28 Jun 2000 over ground KM

volatilis Ma Puu Keokea 7100 ft 2 20 May 2001 over ground KM

volcanicus H Mauna Loa Rd. 1850 m 2 5 Apr 1989 PG

volcanicus H Mauna Loa Rd. 1850 m 1 9 Jul 1989 PG

volcanicus H below 3-Tree Kipuka 1890 m 1 9 Jul 1989 Metrosideros polymorpha PG

volcanicus H Kipuka Maunaiu 1890 m 1 2 17 Jul 1989 PG

volcanicus H below 3-Tree Kipuka 1890 m 2 1 17 Jul 1989 Metrosideros polymorpha PG



volcanicus H south of Bird Park 1230 m 2 4 Aug 1989 PG

volcanicus H Footprints Trail 900 m 1 30 Dec 1989 PG

volcanicus H Kipuka Maunaiu 1890 m 6 31 Dec 1989 PG

volcanicus H Hualalai summit 8000 ft 1 20 Apr 1992 Coprosma sp. SM

volcanicus H Mauna Loa Rd. 2050 m 2 5 21 Mar 1993 JP



volcanicus H Mauna Loa Rd. 4200 ft 1 2 Jan 1999 over ground KM

volcanicus H 0.9 mi. S Mauna Loa Rd. 6000 ft 4 4 Jan 1999 over ground KM

volcanicus H Devastation Trail 3700 ft 3 8 Jan 1999 Styphelia tameiameiae KM

volcanicus H Kipuka Nene 2900 ft 1 1 13 Jan 1999 yellow pan trap KM

volcanicus H Kulani Cone 5100 ft 2 14 Jan 1999 boggy area KM

volcanicus H Kilauea Iki 3900 ft 1 5 Jun 1999 over ground KM

volcanicus H Kilauea Iki 3900 ft 1 8 Jun 1999 over ground KM

volcanicus H Kipuka Nene 2900 ft 1 13 Aug 1999 over ground KM

volcanicus H near Tree Molds 4000 ft 1 11 Jun 2000 over ground KM

volcanicus H Kipuka Puaulu 3900 ft 1 24 Jul 2000 Myoporum sandwicense KM

volcanicus H Crater Rim Trail 4100 ft 2 24 Jul 2000 Dodonaea viscosa KM

volcanicus H Byron’s Ledge 3800 ft 2 29 Jul 2000 Styphelia tameiameiae KM

volcanicus H Puu Waawaa 6200 ft 1 30 May 2001 Dodonaea viscosa KM

volcanicus H Puu Waawaa 6200 ft 4 3 30 May 2001 over ground KM

volcanicus H Napau Trail 3000 ft 1 14 Jul 2002 Metrosideros polymorpha KM

volcanicus Ma Holua Cabin 7200 ft 1 6 Jun 1985 Styphelia tameiameiae CF

volcanicus Ma Haleakala Crater 7600 ft 1 25 Aug 1986 Dubautia Argyroxiphium AM

volcanicus Ma Halemauu Trail 4 27 May 1992 RT

volcanicus Ma Lihau summit 4000 ft 1 8 May 1993 DP

volcanicus Ma base of Puu Maile 2195 m 1 25 Jun 1996 RT

volcanicus Ma Mamane Hill 2255 m 1 1 28 Jun 1996 Styphelia tameiameiae RT

volcanicus Ma near Oili Puu 2050 m 1 1 8 Jul 1996 RT

volcanicus Ma Halemauu Trail 2440 m 1 28 Aug 1996 RT

volcanicus Ma near HALE visitor center 9750 ft 3 11 Sep 1996 Dubautia menziesii RT

volcanicus Ma Upper Koolau Gap 2 11 Sep 1996 RT

volcanicus Ma above Halemauu Trail 1 25 Sep 1996 RT

volcanicus Ma Sliding Sands Trail 2500 m 3 27 Oct 1996 RT

volcanicus Ma Halemauu Trail 2250 m 1 3 Mar 1997 RT

volcanicus Ma Waikane 2010 m 1 8 Apr 1997 RT

volcanicus Ma Lauulu Trail 1955 m 2 2 27 May 1997 RT

volcanicus Ma Sliding Sands Trail 2500 m 3 18 Jun 1997 Sophora chrysophylla RT

volcanicus Ma Kuiki 2170 m 1 7 Aug 1997 RT

volcanicus Ma Science City 3010 m 2 13 Aug 1997 RT

volcanicus Ma Laie flats 2060 m 1 19 Aug 1997 RT

volcanicus Ma Kaupo Trail 1950 m 4 20 Aug 1997 RT

volcanicus Ma Crystal Cave 2285 m 1 25 Sep 1997 RT

volcanicus Ma Paliku 1950 m 1 25 Sep 1997 RT

volcanicus Ma Lauulu Trail 2015 m 1 14 Jan 1998 RT

volcanicus Ma Oili Puu 2020 m 1 15 Jan 1998 Styphelia tameiameiae RT

volcanicus Ma Kipahulu Valley 1980 m 2 21 May 1998 RT

volcanicus Ma Paliku 1950 m 1 4 Nov 1998 RT

volcanicus Ma Sliding Sands Trail 2500 m 1 4 Nov 1998 Dubautia menziesii RT

volcanicus Ma Kuiki 2250 m 2 17 Jan 1999 RT

volcanicus Ma Halemauu Trail 8000 ft 1 12 May 1999 over ground RT

volcanicus Ma Kilohana 2800 m 1 1 Jun 1999 RT

volcanicus Ma Halemauu Trail 7800 ft 1 23 Jun 1999 over ground KM

volcanicus Ma Holua Cabin 2155 m 1 1 Oct 1999 RT

volcanicus Ma Halemauu Trail 7900 ft 1 25 Jun 2000 over ground KM

volcanicus Ma Lauula Trail 8000 ft 2 27 Jun 2000 over ground KM

volcanicus Ma Kalapawili Ridge 7400 ft 1 27 Jun 2000 over ground KM

volcanicus Ma Kaupo Trail 5500 ft 2 28 Jun 2000 over ground KM

volcanicus Ma Halemauu Trail 7900 ft 1 1 29 Jun 2000 Styphelia tameiameiae KM

volcanicus Ma Laie flats 2040 m 1 26 Jul 2000 RT

volcanicus Ma Puu Keokea 7100 ft 1 20 May 2001 over ground KM

sp. A Mo West Kawela Gulch 3700 ft 1 2 Jun 2001 Scaevola chamissoniana KM


INDEX

The index is divided in four parts: Hylaeus species-group names; Arthropod enemies andassociates; Plants used as nesting substrates; and Flower visitation records. Species-groupnames in bold face indicate valid taxa revised in this volume and the pages for their taxonomictreatments. Exotic plants are indicated with an asterisk (*).

Hylaeus species-group names

akoko Magnacca and Daly, 8, 25, 31, 43, 77albonitens (Cockerell), 12andrenoides (Perkins), 8, 26, 37, 45angustulus (Perkins), 8, 29, 41, 47, 78, 80anomalus (Perkins), 7, 8, 31, 38, 50, 100, 106, 184anthracinus (F. Smith), 6, 7, 24, 27, 32, 42, 52, 87,

89, 125assimulans (Perkins), 8, 34, 42, 55, 106, 158, 160assimulans oahuensis (Perkins), 55attripennis (nomen dubium), 209avara Warncke, 198binominatus Meade-Waldo, 198bisinuatus Forster, 12blackburni (F. Smith), 90caeruleipennis (Perkins), 94chlorostictus (Perkins), 8, 28, 38, 58, 81, 89, 106,

184, 187comes (Perkins), 61coniceps (Blackburn), 8, 30, 33, 40, 42, 61, 65, 80connectens (Perkins), 8, 30, 32, 38, 40, 64, 77, 89,

145, 187crabronoides (Perkins), 8, 30, 40, 68, 190, 192difficilis (Perkins), 8, 18, 24, 27, 41, 55, 61, 64, 71,

83, 106, 117, 130, 132, 157, 192, 197, 204, 205,207

dimidiatus (Perkins), 30, 41, 44, 75, 125dumetorum (Perkins), 8, 29, 41, 49, 64, 71, 78,

138, 190, 192erythrodemas (Perkins), 160facilis (F. Smith), 6, 7, 8, 24, 28, 41, 59, 80, 184,

187farinosa (Warncke), 155filicum (Perkins), 8, 28, 40, 44, 83, 114, 119, 145finitimus (Perkins), 7, 8, 24, 34, 38, 86flavifrons (Kirby), 32, 37, 53, 55, 87, 106, 187flavipes (F. Smith), 6, 8, 18, 34, 42, 53, 55, 56, 90,

106, 160fuscipennis (F. Smith), 8, 10, 31, 39, 94, 173fuscipennis obscuripes (Perkins), 94fuscipennis swezeyi (Cockerell), 94gliddenae Magnacca and Daly, 7, 8, 25, 31, 50, 98,

160haleakalae (Perkins), 31, 34, 39, 86, 100, 119hilaris (F. Smith), 8, 9, 10, 33, 36, 55, 58, 94, 103,

111, 117, 193, 195, 201, 205

hirsutulus (Perkins), 8, 27, 38, 106, 166, 168homoeochromus (Perkins), 189hostilis (Perkins), 8, 9, 33, 36, 89, 106, 109, 132hula (Perkins), 29, 39, 44, 75, 86, 112inquilina (Perkins), 8, 9, 32, 36, 106, 114, 189insignis (Forster), 145insignis (Perkins), 145insulae Meade-Waldo, 64kauaiensis (Perkins), 29, 36, 117, 148, 201koae (Perkins), 64kokeensis Magnacca and Daly, 29, 36, 119, 122,

145kona (Blackburn), 8, 29, 39, 75, 77, 122kriechbaumeri Forster, 6kuakea Magnacca and Daly, 24, 25, 27, 32, 125kukui Magnacca and Daly, 8, 31, 39, 127laetus (Perkins), 8, 27, 38, 41, 71, 74, 83, 106, 129,

157laticeps Morawitz, 198laticeps Perkins, 198longiceps (Perkins), 8, 33, 42, 53, 55, 56, 86, 106,

133, 173mana Magnacca and Daly, 8, 24, 27, 29, 41, 135mauiensis (Perkins), 7, 8, 25, 30, 138melanothrix (Perkins), 7, 8, 29, 40, 140mimicus Magnacca and Daly, 8, 28, 39, 84, 86,

119, 138, 142muranus (Warncke), 8, 26, 39, 129, 145, 179mutatus (Perkins), 27, 37, 117, 148nalo Magnacca and Daly, 7, 24, 25, 26, 32, 150neglectus (Perkins), 100niloticus (Warncke), 7, 8, 34, 42, 58, 94, 152nivalis Morawitz, 155nivalis Perkins, 155nivicola Meade-Waldo, 8, 11, 26, 41, 74, 106, 155,

197, 204, 205, 207obscuratus (Perkins), 152obscuratus Schenck, 152ombrias (Perkins), 8, 34, 42, 56, 58, 93, 158ovatula (Warncke), 176paradoxicus (Perkins), 8, 10, 25, 31, 38, 98, 100,

106, 160pectoralis Forster, 6pele (Perkins), 8, 30, 40, 75, 77, 163, 208perkinsianus (Timberlake), 7, 8, 24, 27, 38, 107,

166perspicuus (Perkins), 7, 8, 30, 37, 168


psammobius (Perkins), 24, 26, 31, 42, 152, 170pubescens (Perkins), 8, 9, 10, 11, 25, 31, 39, 44,

94, 97, 173rubrocaudatus (nomen dubium), 209rugiventris Blackburn, 52rugosulus (Cockerell), 179rugulosus (Perkins), 32, 42, 176satelles (Blackburn), 7, 8, 26, 34, 39, 145, 147,

179, 198setosifrons (Perkins), 8, 31, 38, 181simplex (Perkins), 7, 8, 24, 28, 41, 59, 81, 83, 184solaris Magnacca and Daly, 33, 38, 55, 89, 133,

135, 187species A, 8, 40, 208specularis (Perkins), 8, 10, 30, 37, 41, 68, 70, 80,

138, 148, 189sphecodoides (Perkins), 8, 9, 33, 36, 75, 106, 116,

192takumiae Magnacca and Daly, 26, 41, 152, 195,

204trigona Warncke, 64unicus (Perkins), 29, 40, 103, 119, 145, 198vicina Perkins, 64vicinus Sichel, 64volatilis (F. Smith), 8, 9, 10, 33, 36, 58, 75, 83, 106,

116, 195, 197, 201volcanicus (Perkins), 8, 26, 41, 74, 106, 117, 157,

197, 204, 205

Arthropod enemies and associates

Afrocypholaelaps africana (Ameroseiidae), host:H. pubescens, 10, 176

Anoplolepis longipes (Formicidae), 10cleptoparasites, 9Coelopencyrtus kaalae (Encyrtidae), hosts:

H. connectens, 68H. fuscipennis, 98H. pubescens, 176

Coelopencyrtus sexramosus (Encyrtidae), host:H. pubescens, 176

Eupelmus sp. (Eupelmidae), hosts:H. fuscipennis, 98H. setosifrons, 184

Eupelmus nr. E. euprepes (Eupelmidae), host:H. unicus, 201

Iridomyrmex humilis (Formicidae), 10Linepithema humile (Formicidae), 10Nesencyrtus kaalae (Encyrtidae), hosts:

H. connectens, 68H. fuscipennis, 98H. pubescens, 176

Pheidole megacephala (Formicidae), 10Pheidole fervens (Formicidae), 10Vespula pensylvanica (Vespidae), 10

Plants used as nesting substrates

Acacia koaH. pubescens, 176

Boehmeria grandisH. anomalus, 52

Broussaisia argutaH. facilis, 83

Clermontia grandifloraH. facilis, 83

Leptospermum scoparium*H. pubescens, 176

Metrosideros polymorphaH. fuscipennis, 98H. melanothrix, 140H. pubescens, 176

Myrsine lessertianaH. gliddenae, 100

Pipturus sp.H. anomalus, 52H. unicus, 201

Psychotria sp.H. melanothrix, 140

Rubus hawaiiensisH. haleakalae, 103

Smilax sandwicensisH. facilis, 83

Sophora chrysophyllaH. paradoxicus, 160

Tetraplasandra sp.H. kauaiensis, 119

Flower visitation records

Acacia confusa*H. fuscipennis, 97

Acacia koa, 11H. anomalus, 52H. chlorostictus, 61H. connectens, 67H. difficilis, 74H. kokeensis, 122H. laetus, 132H. mimicus, 145H. unicus, 200

Alphitonia ponderosaH. dumetorum, 80

Antidesma platyphyllaH. mutatus, 150

Argemone glaucaH. anthracinus, 55

Argyroxiphium sandwicenseH. difficilis, 74H. nivicola, 157

Bacopa monnieriH. psammobius, 173


Bidens menziesiiH. difficilis, 74H. kona, 125H. laetus, 132H. ombrias, 160H. pele, 166

Bidens sandvicensisH. laetus, 132

Bidens tortaH. connectens, 67

Broussaisia argutaH. connectens, 67

Buddleia asiatica*H. dumetorum, 80

Canavalia galeataH. connectens, 67

Chamaesyce atrococcaH. andrenoides, 47H. hirsutulus, 109H. kokeensis, 122H. laetus, 132H. mutatus, 150

Chamaesyce celastroides, 11H. anthracinus, 55H. difficilis, 74

Chamaesyce degeneriH. anthracinus, 55H. longiceps, 135

Chamaesyce multiformisH. connectens, 67H. laetus, 132

Chamaesyce olowaluana, 11H. akoko, 44H. coniceps, 63H. connectens, 67H. difficilis, 74H. dimidiatus, 77H. filicum, 86H. hula, 114H. kona, 125H. laetus, 132H. ombrias, 160H. paradoxicus, 163H. pele, 166

Chamaesyce spp., 11Cheirodendron trigynum, 11

H. andrenoides, 47H. crabronoides, 70H. difficilis, 74H. dumetorum, 80H. fuscipennis, 97H. haleakalae, 103H. muranus, 147H. pubescens, 176H. setosifrons, 184H. specularis, 192H. unicus, 200

Claoxylon sandwicense, 11H. andrenoides, 47H. chlorostictus, 61H. connectens, 67H. laetus, 132H. mutatus, 150H. specularis, 192

Coprosma montanaH. nivicola, 157H. takumiae, 197

Coprosma sp.H. volcanicus, 207

Crocosmia × crocosmiiflora*H. coniceps, 63H. dumetorum, 80H. pele, 166

Dodonaea viscosa, 11H. difficilis, 74H. flavipes, 92H. laetus, 132H. nivicola, 157H. volcanicus, 207

Dubautia ciliolataH. difficilis, 74

Dubautia laxaH. unicus, 200

Dubautia menziesiiH. difficilis, 74H. nivicola, 157H. volcanicus, 207

Dubautia menziesii × Argyroxiphium sandwicenseH. volcanicus, 207

Dubautia scabraH. difficilis, 74H. dumetorum, 80

Fragaria vesca*H. dumetorum, 80

Geranium cuneatumH. coniceps, 63H. nivicola, 157

Hedyotis terminalisH. dumetorum, 80

Hypochoeris radicata*H. pubescens, 176

Ilex anomalaH. dumetorum, 80H. mimicus, 145

Jacquemontia ovalifolia, 11H. flavipes, 92

Lipochaeta lobataH. assimulans, 58

Melicope sp.H. angustulus, 49H. sp. A, 208

Metrosideros polymorpha, 11H. andrenoides, 47H. coniceps, 63


Psychotria sp.H. anomalus, 52

Rubus penetrans*H. angustulus, 49

Rubus rosifolius*H. connectens, 67H. dumetorum, 80

Rubus sp.H. setosifrons, 184

Santalum ellipticumH. longiceps, 135

Santalum freycinetianumH. laetus, 132H. mana, 138

Santalum paniculatumH. coniceps, 63H. difficilis, 74H. hula, 114H. laetus, 132H. pubescens, 176H. setosifrons, 184

Sapindus saponariaH. difficilis, 74

Scaevola chamissonianaH. coniceps, 63H. connectens, 67H. haleakalae, 103H. unicus, 200H. sp. A, 208

Scaevola coriaceaH. longiceps, 135

Scaevola gaudichaudianaH. connectens, 67H. mimicus, 145H. unicus, 200

Scaevola proceraH. connectens, 67H. kauaiensis, 119H. laetus, 132H. specularis, 192

Scaevola sericeaH. anthracinus, 55H. difficilis, 74H. flavifrons, 89H. flavipes, 92H. hostilis, 111H. longiceps, 135H. ombrias, 160H. solaris, 189

Scaevola spp., 11Schinus terebithifolius*

H. difficilis, 74Sesbania tomentosa, 11

H. anthracinus, 55H. flavipes, 92H. laetus, 132H. longiceps, 135

H. connectens, 67H. difficilis, 74H. dumetorum, 80H. fuscipennis, 97H. haleakalae, 103H. hirsutulus, 109H. kauaiensis, 119H. laetus, 132H. mimicus, 145H. muranus, 147H. pubescens, 176H. rugulosus, 179H. setosifrons, 184H. specularis, 192H. unicus, 200H. volcanicus, 207

Myoporum sandwicense, 11H. anthracinus, 55H. chlorostictus, 61H. coniceps, 63H. connectens, 67H. crabronoides, 70H. difficilis, 74H. dumetorum, 80H. filicum, 86H. flavipes, 92H. hula, 114H. inquilina, 116H. laetus, 132H. longiceps, 135H. paradoxicus, 163H. pele, 166H. pubescens, 176H. specularis, 192H. volcanicus, 207

Myrica faya*H. dumetorum, 80

Oenothera sp.*H. nivicola, 157

Pisonia brunonianaH. dumetorum, 80

Plantago lanceolata*H. laetus, 132

Portulaca oleraceae*H. flavifrons, 89

Psychotria faurieiH. unicus, 200

Psychotria hawaiiensisH. dumetorum, 80H. pubescens, 176H. setosifrons, 184

Psychotria marinianaH. andrenoides, 47H. connectens, 67H. mimicus, 145

Psychotria mauiensisH. fuscipennis, 97


Sesuvium portulacastrumH. psammobius, 173

Sida fallax, 11H. anthracinus, 55H. assimulans, 58H. flavifrons, 89H. flavipes, 92H. laetus, 132H. longiceps, 135H. ombrias, 160H. perkinsianus, 168H. solaris, 189

Sophora chrysophylla, 11H. coniceps, 63H. connectens, 67H. difficilis, 74H. flavipes, 92H. kona, 125H. laetus, 132H. nivicola, 157H. pele, 166H. volcanicus, 207

Styphelia tameiameiae, 11H. chlorostictus, 61H. connectens, 67H. difficilis, 74

H. dumetorum, 80H. flavipes, 92H. hirsutulus, 109H. hostilis, 111H. inquilina, 116H. kauaiensis, 119H. laetus, 132H. mutatus, 150H. nivicola, 157H. takumiae, 197H. volatilis, 204H. volcanicus, 207

Tetraplasandra oahuensisH. fuscipennis, 97H. unicus, 200

Tibouchina urvilleana*H. pubescens, 176

Tournefortia argentea*, 11H. anthracinus, 55H. difficilis, 74H. flavipes, 92

Tribulus cistoidesH. flavipes, 92H. ombrias, 160

Vaccinium reticulatumH. difficilis, 74

Vitex rotundifoliaH. longiceps, 135

PREVIOUS VOLUMES OF INSECTS OF HAWAII

I"':"'Reprint of 19 48 INTRODUCTION. Elwood C. Zimmerman , with n ew Foreword by J ames K. Li ebherr and sh ortbiography of au th o r by J ames O . Juvik. 256 pages, illustra ted . 2001.Geological Hi story of H awaii: Derivation , Disp ersal, and Di stribution , Evolution and Devel opment, Analysesand Summaries of Biota.

16-HAWAIIAN CARABIDAE (Co LEOPTERA) .James K. Liebherr and Elwood C. Zimmerman . 504 pages, illustrated. 2000.Part 1: Introdu ction and Tribe Platynini

17-HAWAIIAN H t'LlEUS (NESOPROSOPlS) BEES. Howell V. Daly and Karl N. Magn acca . 240 pages, illustrated. 2003.Hymen op te ra: Apoi dea

NO LON GER AVAILABLE

I-INTRODUCTION. Elwood C. Zimm erman . xx + 206 pages; 52 cuts. 1948.Geological Histor y of Hawaii; Derivation, Dispersal and Distribution, Evolution and Development, Analyses andSummaries of th e Biota. [Reprin t available, see above.]

2-APTERYGOTA TO THYSANOPTERA. Elwood C. Zimm erman . x + 475 pages; 228 cuts . 1948.. Th ysanura, Dip lura, Protura, Coliembola, Orthoptera, Isoptera , Embioptera, Dermaptera, Zoraptera , Corrodentia ,

Mallophaga, Anop lura, Odon ata, Th ysanoptera.

3- HETEROPTERA. Elwood C. Zimmerman . x + 225 pages; 110 cuts. 1948.Cydnidae, Pentatomidae, Coreidae, Lygaeidae, Tingidae , Eni coc ephalidae , Reduviidae , Nabidae, Cimicidae,Anthocoridae, Cr yptosternmatidae, Miridae , Saldidae, Hebridae, Mesovel iidae, Veliid ae, Ger ridae , Not on ectidae,Cori xidae.

4-HOMOPTERA: AUCHENORHYNCIJA. Elwood C. Zimmerman. x + 268 pag es; 92 cuts . 1948.Cercopidae, Cicadellidae, Membracidae , Cixiidae, Delphacidae, Flatidae .

5- HoMOPTERA: STERNORHYNCHA. Elwood C. Zimm erman. x + 464 pages; 238 cuts. 1948.Psylloide a, AJeyrodo idea, Aph idoidea, Coccoid ea .

6-EpHEMEROPTERA-NEUROPTERA - TRICHOPTERA AN DSUPPLEMENT TO VOLS. 1-5. Elwood C. Zimmerman . ix + 209 pages;105 cuts . 1957.

7-MACROLEPIDOPTERA. Elwood C. Zimm erman . xiv + 542 page s; 425 cuts. 1958.Geometridae, Noctuidae, Sphingidae, Pieridae, Nymphalidae, Danaidae, Lycaen idae.

8- LEPIDOPTERA: PYRALOIDEA. Elwood C. Zimm erman . xii + 456 pages; 347 cuts. 1958.Galleriinae, Pyraustinae, Scop ari inae , Nymphulinae, Pyralinae, Crambinae , Ph ycitinae , Pterophoridae, Alucitidae.

9-MICROLEPIDOPTERA. Elwood C. Zimm erman. xxiv + 1903 pag es; 1355 cu ts; 8 color plates. 1978.Monotrysia, Tin eoidea, Tortricoidea, GraciIlarioidea, Yponomeutoidea, AJucitoidea, Gelechioidea.

lO-DIPTERA: NEMATOCERA-BRACHYCERA (EXCEPT DOLlCIJOPODIDAE). D. Elmo Hardy. xii + 368 pages; 120 cuts . 1960.Tipulidae, Psychodidae, Culicidae, Ch ironomidae, Ceratopogonidae, Scatopsidae, Mycetophilidae, Sciaridae,Cecidomyiidae , Stratiomyidae , Bombyliidae , Sce nopinidae , Empididae .

11- DIPTERA: BRACHYCERA II - CYCLORRHAPHA 1. D. Elmo Hardy. x + 458 pages; 140 cuts. 1964.Dolichopodidae, Phoridae , Lon ch opteridae, Pipunculidae, Syrphidae.

lI-SUPPLEMENT-DIPTERA: DOLICHOPODJDAE AND APPENDIX (PHORIDAE) .JoAnn M. Tenorio. vi + 73 pages; 7 cuts. 1969.

12- DIPTERA: C YC LORRHAPHA II. D. Elmo Hardy. x + 814 pages; 352 cu ts. 1965.Series Sch izophora, Section Acalypterae I, Famil y Drosophilidae.

l3-DIPTERA: CYCLORRHAPHA III. D. Elmo Hardy and M. D. Delfinado. vi + 451 pages; 179 cuts. 1980.Series Sch izoph ora, Section Acalypterae, Exclus ive of Fam ily Drosophilidae .

14-DIPTERA: CYCLORRHAPHA IV. D. Elmo Ha rdy. viii + 491 pages; 198 cuts. 1981.Series Schizophora, Section Calyptratae.

15-CoLLEMBOLA. Kenneth Christian sen and Peter Bellin ger. viii + 446 pages; 134 figu res , 162 plates , 99 map s. 1992.Suborder Arthropleona; Sub order Symphypleona.

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